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SOUTH 

AUSTRALIAN 
MUSEUM 



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VI AW 



NEW SPECIES AND NEW RECORDS OF CLOACINA VON LISTOW, 1898 

(NEMATODA: STRONGYLOIDEA) PARASITIC IN MACROPODID 

MARSUPIALS FROM PAPUA NEW GUINEA 

/. Beveridge 



Summary 

New species described from macropodid marsupials in Papua New Guinea are : Cloacina cretheis 
sp. nov. from tree kangaroos, Dendrolagus inustus (type host), D. dorianus, D. goodfellowi, D. 
matchiei and D. scottae; C. cunctabunda sp. nov. from D. mbaiso; C. eurynome sp. nov. from D. 
dorianus and D. scottae; C. hecale sp. nov. from D. dorianus; C. theope sp. nov. from D. matschiei 
and D. dorianus; C. erigone sp. nov., C. hyperaea sp. nov., C. nephele sp. nov., C. polymena sp. 
nov., C. praxithea sp. nov. and C. procris sp. nov. from the scrub wallaby, Macropus agilis. New 
host records are : C. caballeroi Mawson, 1977, C. sterope Beveridge & Speare, 1999 and C. syphax 
Beveridge & Speare, 1999 in Do. Hageni. Additional geographical records are given for C. cornuta 
in M. agilis. A key to the known species of Cloacina in Papua New Guinea is provided. 



NEW SPECIES AND NEW RECORDS OF CLOACINA VON LINSTOW, 1898 

(NEMATODA: STRONGYLOIDEA) PARASITIC IN MACROPODID 

MARSUPIALS FROM PAPUA NEW GUINEA 



t. BEVERIDGE 



BEVERIDGE. 1. 2002. New .species and new records of Cloacina von Linsiow. 1898 
iNematoda: Sirongyloidca) parasitic in maeropodid marsupials from Papua New Guinea. 
Records of the South Australian Museum 35(1): I -32 

New species described from maeropodid marsupials in Papua New Guinea are : Cloacina 
creiheis sp. nov from tree kangaroos, Dcndndagus inustus (type host), 0. dorumus, D. 
got'ilfellowi, 0. nmischiei and 0. scoriae; C. cunctabunda sp. nov from 0. mbaiso; C. 
emynome sp. nov from 0, darianus and 0. scotlae; C. hecale sp. nov. from 0. dorianus; C 
iheope sp. nov from 0. matschiei and 0. darianus; C. erigone sp. nov.. C. hyperaea sp. nov , 
C. nephete sp. nov.. C polymelia sp. nov., C. praxillwa sp. nov. and C. procris sp. nov. from 
the scrub wallaby. Diimipxis hageni. and C. oweni n. sp. and C. papuensis ti. sp. from the 
agile wallaby. Matropus agilis. New host records arc: C cahalleroi Mawson, 1977, C. sleropc 
Beveridge & Speare. 1999 and C, syphax Beveridge & Speare, 1999 in Do. hageni. Additional 
geographical records are given for C. Camilla in M. agilis. A key lo the known species of 
Vtoctointl in Papua New Guinea is provided. 

I Beveridge, Department of Veterinary Science. University of Melbourne, Parkville. Victoria 
.1052. Australia. Manuscript teceived 7 May 2001. 



Species of the nematode genus Cloacina von 
Linstow. i 898 occur exclusively in the stomachs 
of maeropodid marsupials. Currently, 103 species 
are recognised as valid (Beveridge 1998, 199% 
Beveridge and Speare 1999) although substantial 
numbers of species remain to be described 
(Beveridge 1998). Most of the species described 
to date are Tram Australian kangaroos and 
wallabies, with relatively few records from Papua 
New Guinea. This situation reflects the paucity of 
knowledge Of the parasite fauna of macropodids 
from Papua New Guinea, which is based currently 
on a small number of incidental collections 
Nevertheless, material currently available from 
various kangaroos and wallabies consists of a 
number of novel species of Cloacina. Thirteen 
new species are described in the current paper as 
well as new host and distribution records. The 
opportunity is taken to provide a summary of 
species of Cloacina known from Papua New 
(]umea together with a key to facilitate their 
identification Finally, a preliminary comparison 
is made between the Australian and Papua New 
Guincan species and their host distributions. 

Beveridge (1998) noled the occurrence of a 
number of species of Cloacina in Papua New 
Guinea, specifically C. austral is (Yorke & 
Maplcstonc, 1926) in the agile wallaby, Macropus 
agilis (Gould. 1842); C. cahalleroi Mawson, 1977 



in the grey scrub wallaby. Dorcopsis luctuosu 
(D'Albertis, 1874) and the brown scrub wallaby. 
D muelleri (Schlegel, 1866); C cloelia Beveridge. 
1998 in the pademelons Thylogale calabyi 
Flannery, 1992 and T. slignuuica (Gould, I860). 
C. coniuta (Davey & Wood, 1938) in M. agilis. 
C. tybele Beveridge. 1998 in T. stigmatica; and 
C. dahli von Linstow, 1898 in Thylogale brownt 
Ramsay, 1877, T. calabyi and T. stigmatica. 
Subsequently, based on an examination of the 
parasites of four small scrub wallabies, 
Dorcopsulus vanheiirni (Thomas, 1922). 
Beveridge ci Speare (1999) described seven new 
species, C. sancus, C. sciron, C. sappho. C. solan. 
C. solymus, C. stempe and C. syphax Beveridge. 
(1998) noted the presence of undescribed species 
from the white-striped scrub wallaby, Dorcopsis 
hageni Heller, 1897 in the collections of the South 
Australian Museum, Adelaide, and from Da. 
luctuosu in the collections of The Natural History 
Museum, London. Flannery el al. (1996) also 
reported species of the genus Cloacina in a vartely 
of tree kangaroos. Dendrolagus inustus Mueller, 
1 840. D. goodfcllowi Thomas, 1 908, D. dorianus 
Ramsay. 1883; D. scomw Flannery & Seri, 1990; 
and D. mbaiso Flannery, Boeadi & Szalay, 1995. 
a group of kangaroos not previously known to 
harbour Cloacina. These various undescribed 
collections form the basis of the current report. 



I. BEVERIDOfc' 



Materials And MirrnoDS 

Nematodes were examined from collections 
held in the Australian Helminthologieal Collection 
(AHC), South Australian Museum (SAM'), 
Adelaide, the Natural History Museum (BMNH), 
London and the Unilcd States National Parasite 
Collection (USNPC), Bcltsville. Maryland. 
Nematodes were washed in water and cleared in 
lactophenol. Drawings were made using a drawing 
tube attached to an Olympus BII2 microscope, 
Drawings of apical views of the heads of 
nematodes arc oriented with the dorsal aspect 
uppermost; drawings of the bursa have the ventral 
surface uppermost, All drawings are of paratype 
specimens. Measurements were made with an 
ocular micrometer. All measurements are in 
millimetres and are presented as the range 
followed by the mean in parentheses. 

Morphological terminology follows Beveridge 
(1998). Because of the relative uniformity of the 
genus Cloacina, the descriptions presented are 
abbreviated and follow the format used by 
Beveridge (1998), in which a full description of 
the genus was followed by individual species 
descriptions concentrating on features of 
diagnostic significance. A detailed description of 
the genus is available in Beveridge (199S). 
Features which arc relatively invariable, such as 
the disposition of ventral and lateral bursal rays, 
are not included in descriptions but are. 
nonetheless, illustrated. 

Types of the new species have been deposited 
in SAM, BMNH and USNPC. Host nomenclature 
follows Flatmery (1995) and Flannery el a I. 
(1995). 

Following previous practice (Beveridge 1998"), 
the names of new .species are based on those of 
classical deities since Cloacina was the Roman 
goddess of the toilet. An invocational quatrain on 
this subject, usually attributed to Byron (see 
Lewin 1999, p. 31), occurs in a number of 
variations, but the rendering by Lewin (1999) is 
presented here; 

O Cloacina. goddess of this place. 
Look on thy suppliants with smiling face, 
Soft yel cohesive let the ir offering-; Hnw, 
Noi rashly swift nor insolently slow 

Sysi-EMATICS 

Cloacina caballeroi Mawson, 1977 
Material examined: 16, 13 5, from stomach of 



Dorcopsis hageni, Usino, Madang, Papua New 
Guinea, coll. f. Reardon, May 1987, SAM AHC 
31425. 

Remarks 

Cloacina caballeroi was described by Mawson 
(1977) and Beveridge (1998) from Dorcopsis 
muellen and Do, luctuosa from Irian Jaya and 
Papua New Guinea. This represents the first 
record from Do. hageni 



Cloacina cornuta (Davey & Wood, 1938) 

Material examined: 26, 15, from slornuch of 
Macropus agilis, Bula Plain, Bensbach, Papua 
New Guinea, coll. I. Owen, May, 1998, BMNH 
1998.9.28.24-26. 

Remarks 

Claaciiw cornula was reported from Macropus 
agilis from Port Moresby and Cape Rodney by 
Beveridge (1998), The present record adds an 
additional locality. Data provided by Dr I. Owen 
indicate that in four wallabies examined, numbers 
of C. cornuta ranged from 1650-9600 (mean 
4700). Although based on a very small number of 
wallabies, these data contrast strikingly with those 
of Spearc et al. (1983) for M. agilis from northern 
Australia in which C. cornuta was found in only 
41% of hosts examined, and with Beveridge et al. 
(1998) who found the nematode in 35% of 
wallabies in Queensland. In Australia, C. cornula 
invariably occurs in much lower numbers than the 
synhospitalic species C. australis (unpublished 
observations). 



Cloacina cretheis sp, nov. 
(Figs 1-13) 

Types; From stomach of Dendrolagus imtstus. 
Mt Somoro, Sandaun Province. Papua New 
Guinea, coll. T. Flannerv. 10iii.l990. Holotvpe 
o\ SAM AHC 31426: allotype 9. SAM AHC 
31427. Paratypes: I2d, 125, SAM AHC 31428; 
Id. I 9, BMNH 2001.4.10.1-2; IeT, I 9, USNPC 
91 133. Slide preparations of male, apical views of 
mouth and bursa, SAM AHC 28378. 

Material examined: From Dendrolagus imtstus: 
types. From Dendrolagus dotictntis: I o* , 4 9, 
Gunung Ki, Tembagapura. Irian Jaya. coll. T. 
Flannery, I9.V.1994. SAM AHC 31430; 26. 109, 
Lake Trist, Papua New Guinea, coll. 1. Redmond, 
UMNH, 1979, 4486-4505. From Dendrolagus 



CL0AC1NA FROM PAPUA NEW GUINEA 




FIGURES 1-13. Cloacma crciheh sp. nov., specimens from Dendrolagus inastuf. I. Anterior end, lateral view. 2. 
Cephalic extremity, lateral view, dorsal aspect on right-hand side 3. Cephalic extremity, dorsal view. 4. Submedian 
cephalic papilla. 5. Cephalic extremity, apical view. 6. Cephalic extremity, transverse optical section through 
buccal capsule. 7. Genital cone, ventral view. 8. Genital cone, dorsal view. 9. Gubemaeulum, ventral view. 10. 
Bursa, apical view. 11. Distal tip of spicule, lateral view. 12. Female tail, lateral view, 13. Female genital system, 
lateral view. Scale bars: 1, 10, 12. 13, 0.1 mm; 2-9, 1 1, 0.01 mm. 



I Kl'VERIDGE 



vomlfelluwi. ajj, 59. Mt Machold. Sandaun 
Province, Papua New Guinea, coll. T. Flannerv 
and K. Vula, H.xii.1990. SAM AHC 3143L 
From Dcndrolagus maischiei Forster & 
Rothschild, 1907; 26(3. 28 V. Huon Peninsula, 
Papua New Guinea, coll. J. Mayer. July 1999, 
USNPC 91146, From Dendmla^nix scottae; 9o*. 
52, Sweipini, Sandaun Province, Papua New 
Guinea, coll T Flannery. 15. vi. 1991, SAM AHC 
31429, 

Description 

Small nematodes; cervical cuticle very 
slightly inflated to beyond level of excretory 
pore; transverse cuticular annulations widely 
spaced, 0.030 apart. Submedian cephalic 
papillae prominent. 0.015 long, projecting 
anteriorly from peri-oral cuticle; distal segment 
sub-spherical 0.005 long, shorter than 
cylindrical proximal segment, 0.0IQ long, 
Buccal capsule shallow, symmetrical in dor.sai 
and lateral views, circular in transverse section, 
wider than deep, wall without striations. Leaf 
crown elements 8 in number, slightly recurved 
at tips; peri-oral cuticle not inflated into lobes 
attached to each element. Dorsal sector of 
oesophagus hearing opening of dorsal 
oesophageal gland, not projecting into lumen of 
buccal capsule. Oesophagus, slender, clavifoim. 
slightly wider at posterior end; lining without 
selcrouscd busses, denticles absent. Nerve ring 
in mni-oesupliageal region; deirids posterior to 
nerve ring; excretory pore at level of 
ocsophago-intcstinal junction, 

Male. Measurements of 5 specimens, types. 
Total lenglh 6.7-9.0 (7 8); maximum width 0.35- 
045 (0.40); buccal capsule 0.010 (0.010) x 0.023- 
0.025 (0.023); oesophagus 46-0.57 (0.52): 
nerve ring to anterior end 0.20-0.24 (0.22), 
excretory pore to anterior end 0,39-0.48 (0.451. 
deirid to anterior end 0.28-0.36 (0.32), spicules 
2.M- 2.62 (2.541; gubernaculum 0.03 (0.03) long. 

Measurements of 5 specimens from D. 
gtjodfeilowi: total length 7.9-9.3 (S.8); maximum 
width 0.48-0.56 (0.51); buccal capsule 0.010- 
012 (0.011) x 0O23-O.025 (0.023); oesophagus 
0.53-0.64 (0.59); nerve ring to anterior end 0.23- 
0.24 (0 24). excretory pore to anterior end 0.44- 
0.53 (0 471. deirid to anterior end 0.20-0.34 
(0.29,; spicules 2.53-2,94 (2.74). gubernaculum 
0.030-0 040 (0.034) long. 

Measurements ot 3 specimens Irom D, scaiuw: 
total length 6.9-9.7 (8.4), maximum width 0.43- 
0.47 (0 45 1, buccal capsule 0,010 (0.010) x 0.023- 
0.025 (0.023); oesophagus 0.48-0.62 (0.56); 



nerve ring to anterior end 0.20; excretory pore lo 
anterior end 0.40-0.46 (0.42); deirid lo anterior 
end 0.25-0.31 (0.28); spicules 2.63-2.66 Ctoi. 
gubernaculum 0.030-0.035 (0.032) long. 

Measurements of single specimens from £>. 
dorianus. total length 9.4 ; maximum width 0.43; 
buccal capsule 0.010 x 0.025 ; oesophagus 0.50 . 
excretory pore lo anterior end 0.50; deirid to 
anterior end 0.26; spicules 2.97; gubernaculum 
0.035 long. 

Dorsal ray elongate; external branchlets arise 
at 1/3 length, before major bifurcation; angle of 
bifurcation acute, external branchlets as long as 
internals, directed postero-laterally, not 
reaching margin of bursa; internal branchleis 
originate 2/3 along dorsal ray, directed posKio 
laterally, almost reaching margin of buisa. 
Externo-dorsal ray nor reaching margin of 
bursa. Gubernaculum prominent, wider than 
long Spicule tip blunt, ala terminates anterior 
to spicule tip; anterior Up of genital cone 
conical; posterior lip with paired projections 
and cuticular inflation of internal surface of 
bursa on either side 

Femak Measurements of 5 specimens, types. 
Total length 7.7-9.0 (8.2); maximum width 6. 5 2 - 
54 (0.52); buccal capsule 0.008-0.010 (0.010) x 
0.023-O.025 (0.023); oesophagus 48-0 55 
(0.53); nerve ring to anterior end 0.20-0.30 
0>.23); excretory pore to anterior end 0.39-0.50 
(0.44); deirid to'anterior end 0.26-O.33 (0.30); tail 
0.23-0.31 (0.27), vulva to posterior end 35-0.40 
(0.38); vagina 0.50-0.73 (0.62); egg 0.075-0 0§Q 
(0.080) x 0.040-0.050 (0.045). 

Measurements of 5 specimens from D. scviun: 
total length 8.8-10.7 (T0.0); maximum width 
0.49-0 64 (0 59); buccal capsule 0.008-0.01 
(0.009) x O.02O-O.025 (0.023); oesophagus 0.53- 
0.58 (0,55); nerve ring to anterior end 0.20: 
excretory pore to anterior end 0.48-0.50 (0.49); 
dc'rnd lo anterior end 0.29-O 32 (0.30); tail 0.2.3- 
0.32 (0.28); vulva lo posterior end 0.38-0.40 
10.39); vagina 0.53-O.5R<0.56). 

Measurements of 3 specimens from D. 
dorianus: total lenglh K.8-9.2 (9.0); maximum 
width 49-0.72 (0.60); buccal capsule 010- 
0.013 (0.012) x 0.023-0.025 (0.024); oesophagus 
0.53-0 60 (0,57); nerve ring lo anterior end 0~23 
(0.23); excretory pore to anterior end 0.48-0.51 
(0 49); deirid to anterior end 20-0.29 (.0.24); tail 
0.21-0.25 (0.23); vulva to posterior end 0.49-0.50 
(0.49); vagina 0.64-0 67 (0.66); egg 0.070-0.075 
(0.07S) x 6.040-0.050 (0.045). 

Female tail short, conical; vagina elongate, 
convoluted; egg ellipsoidal. 



CWAC1NA FROM PAPUA NEW GUINEA 



Remarks 

C. crelheis sp. nov. is characterised by 
submedian papillae with a small sub-spherical 
distal segment, a simple buccal capsule, eight 
elements to the leaf crown, a slender, 
unornamented oesophagus, deirids posterior lo 
the nerve ring, a dorsal ray in which the 
external branchlets arise before the major 
bifurcation and an elongate, convoluted vagina. 
The shape of the dorsal ray alone separates it 
Irom all congeners except C cabalteioi, C. 
etyo Bcveridgc. 1998. C. ips Beveridgc. |{*9B, 
and C. u'jiliu.x. It differs from all of these 
except C labolkroi in having the deirid 
posterior to ihe nerve ring, although in some 
highly contracted specimens of C, crelheis, in 
which the anterior oesophagus becomes 
sinuous, the deirid may appear more anteriorly 
on the contracted cuticle. C. cretheis is furthet 
distinguished from C. syphax which has a 
sinuous anterior margin to its buccal capsule 
and submedian cephalic papillae with enlarged 
distal segments. C. creiluts is distinguished 
Irom C. cubalteroi by spicule lengths, which 
arc 2 .34-2.97 mm in C. cretheis compared with 
I 23-1 45 mm in C. caballeroi. The spicules of 
(' W}/0 1FC 6X-OHH mm long whiK- ihos<" of 
1 ips are 1,36-1.39 mm in length, providing a 
further means of distinguishing the latter 
species. 

C. cretheis occurs in a wide range of tree 
kangaroo species in Papua New Guinea and was 
by far the most frequently encountered species in 
tree kangaroos in this study. It does not occur in 
either Dendrola^us hennettianus DeVis, 1 887 or 
D lumhnltzi Collett, 1S84 in Australia (Spratt et 
al. 1 9'--* I ). Although incomplete, the measurements 
of specimens from different hosts suggest that 
there ate no major differences induced by the host 
species. Some of the minor differences observed 
are due to the state of contraction or of 
preservation of specimens, and (he incomplete sets 
of measurements reflect the poor slate of 
preservation of many of the specimens, precluding 
Hie measurement of internal organs. 



Clour ina cunciabunda sp, nov 

(Figs 14-23) 

Types: From stomach of Dendmln^us inhciso, 
Currip Ridge, Tembagapura. Irian Jaya. col). T. 
Flannery, 24.V.1994. Holotype o\ SAM AHC 
31432; allotype 8, SAM AHC 31433, Paratype 
<J, 3 slides, SAM AHC 28379. 



Description 

Small nematodes; cervical cuticle slightly 
inflated to beyond level of excretory pore; 
transverse cuiicular annulnlions widely spaced, 
0.015 apart. Submcdittti cephalic papillae 
prominent, 0.010 long, projecting anteriorly from 
peri-oral cuticle; distal segment ovoid, 0.006 long, 
longer than cylindrical proximal segment, 0.004 
long. Buccal capsule shallow, .symmetrical in dorsal 
and lateral views, oval in transverse section, 
slightly deeper dorso-vcntrally. wider than deep, 
wall without striations. Leaf crown elements 6 in 
number, recurved at tips; peri-ota! cuticle, inflated, 
inflations not attached to each element of leaf 
crown. Dorsal sector of oesophagus bearing 
opening of dorsal oesophageal gland, not projecting 
into lumen of buccal capsule. Oesophagus slender, 
claviform, slightly wider at posterior end; lining 
without sclerotised tosses; denticles absent Nerve 
ring in mid-oesophageal region, deirids anterior to 
nerve ring, mid-way between nerve ring and buccal 
capsute; excretory pore anterior to level of 
uesophago-inteslinal junction. 

Male. Measurements of holotype and paratype. 
Total length 3.30, 3.32; maximum width U. 16. 
0.17; buccal capsule 0.008, 0.010 x 0.025, 0.025. 
oesophagus 0.34, 0.35; nerve ring to anterior end 
0.1 6. 0.18; excretory pore to anterior end 0.26. 
0.30, deirid io anterior end 0.11, 0.14; spicules 
1.39, 1.57; gubcrnaculum 0.015, 0.015 long. 

Dorsal lobe of bursa and dorsal ray elongate, 
ray broad at origin; external branchlets arise at 1/2 
length, immediately before major bifurcation; 
angle of bifurcation acute; external branchlets 
shorter than internals, directed postcro-latcraUy, 
not reaching margin of bursa; internal branchlets 
originate soon after externals arise from dorsal 
ray, directed postero-laterally, almost reaching 
margin of bursa f?. Memo-dorsal ray not reaching 
margin of bursa. Gubernaculum prominent, wider 
than long. Spicule tip not evened, Anterior lip of 
genital cone conical, posterior lip with paired 
projections. 

Female. Measurements of allotype. Total length 
4.25. maximum width 0.21, buccal capsule 0.010 
x 0.025; oesophagus 0.40; nerve ring to anterior 
end 0.18; excretory pore to anterior end 0.29; 
deirid to anterior end 0.11; tail 0.35; vulva to 
postenor end 0.55; vagina 0.35. 

Female tail elongate, conical; vulva 
immediately anterior to anus, vagina short with 
single convolution; egg not seen. 

Remarks 
Although described from only three specimens. 



I. BEVER1DGE 




FIGURES 14-23. Cloacina cunclabunda sp. nov., specimens from Dendrolagus mbaiso. 14. Anterior end. lateial 
view. IS, Cephalic extremity, lateral view, dorsal aspect on left-hand side. 16. Cephalic extremity, ventral view. 17. 
Submedian cephalic papilla, lateral view. 18. Cephalic extremity, apical view. 19. Cephalic extremity, transverse 
optical section through buccal capsule. 20. Bursa, apical view. 21. Gubcrnaculum, ventral view. 22. Female tail, 
lateral view. 23, Female genital system, lateral view. Scale bars: 22, 23, 0.1 mm; 14-21 0.01 mm. 



CWACINA FROM PAPUA NEW GUINEA 



this species is sufficiently distinctive to allow 
icady recognition, and comes from a host 
restricted to isolated localities in Irian Jaya which 
is unlikely to be Sampled extensively m the future 
(Flannery et al. 1995). Furthermore, this species 
of Cloacina was not found in any other species of 
tree kangaroo 

C cunctabunda is distinguished from all 
congeners except C. cretheis, C, caballerai, C. 
eayo, C. ips and C syphax in possessing a simple 
buccal capsule, an elongate, unornamenled 
oesophagus and the external branchlets of the 
dorsal ray arising before the major bifurcation 
The position of the deirid anterior to the nerve 
ring distinguishes the species from C cretheis and 
C. cabalieroi, while the shape of the buccal 
capsule and the submedian cephalic papillae 
distinguish it from C. syphax. C. cunctabunda is 
distinguished from C. eayo on the basis of spicule 
lengths, which are 1.39-1.57 mm long in C. 
cuiictabioida compared svith 0.68-0.88 mm in C. 
ettyo. C. cuncrabu'ida therefore most closely 
resembles C. ipx. from which it is distinguished 
by the number of leal crown elements (six in C. 
cunctabunda, eight in C ips). in Ihe absence of a 
dorsal oesophageal looili which projects 
pioiuincnlly into the huccal capsule of C ipt\ in 
the shape of the female tail which is slender and 
elongate in C. curutubundu but short and conical 
in C. ips, and in the shape of the vagina which is 
straight in ('. ips but convoluted in C 
runctiibunda 



Chiuiina erigune sp. nov. 

(Figs 24-36) 

Types: From stomach of Dorcopsis hageni, 
Usitio, Madang, Papua New Guinea, coll. T. 
Keardon. May" 1987. Holoiype 6. SAM AHC 
"U4"*4; allotype $> , SAM AHC 31435. Paratypes: 
4jJ, 59. SAM AHC 31436, Id, 19. BMM1 
2001.4. 1 0.3-*. 1d r 1?. USNPC 91134. Slide 
prcparaiions of apical views of mouth and bursa, 
SAM AHC 28380. 

Material examined: types. 

Description 

Small nematodes; cervical cuticle slightly 
inflated to beyond level of excretory pore; 
transverse cuticular angulations widely spaced, 
0.0I5 apart. Submedian cephalic papillae 
prominent, 0.013 long, projecting anteriorly 
from perioral cuticle; distal segment ovoid. 
(1 1)05 long, .shorter than cylindrical proximal 



segment, 0.008 long. Buccal capsule shallow, 
symmetrical in dorsal and lateral views, circular 
in transverse section, wider than deep, wall 
without striaiions. Leaf crown elements 8 in 
number, slightly recurved at lips, margins 
prominently thickened; peri-oral cuticle not 
inflated into lobes attached to each element. 
Dorsal sector of oesophagus bearing opening of 
dorsal oesophageal gland, projecting 
prominently into lumen of buccal capsule. 
Oesophagus claviform, robust, slightly wider at 
posterior end; lining with sclcrolised bosses 
extending from anterior end, halfway to nerve 
ring; single dorsal denticle present in prominent 
preneural swelling of oesophagus. Nerve ring in 
mid-oesophageal region, deirids posterior to 
nerve ring, almost at level of excretory pore; 
excretory pore al level of ocsophago-intestinal 
junction. 

Male. Measurements of 8 specimens, types. 
Total length 2.88-5.11 (4.02); maximum width 
0.18-0.31 (0.26); buccal capsule 0.007-0.010 
(0.008) x 0.023-0.035 (0.032): oesophagus 0.26- 
0.39 (0.34); nerve ring to anterior end 0.14-0.22 
(0.18); excretory pore to anterior end 0.25-O.40 
(0.33); deirid CO anterior end 0.23-0.37 (0.32); 
spicules 1.35-1.79 (1.59); gubemaculum 0.020- 
0.030 (O.024) long. 

Dorsal ray elongate, broad at origin; external 
branchlets arise at 1/2 length, immediately before 
major bifurcation; angle of bifurcation acute; 
external branchlets shorter than internals, directed 
laterally, not reaching margin of bursa; internal 
branchlets originate immediately after externals, 
directed postero-laterally, almost reaching margin 
of bursa. Externo-dorsal ray not reaching margin 
of bursa. Gubemaculum prominent, slightly widt-r 
than long. Spicule tip blunt; ala terminates 
abruptly anterior to spicule tip; anterior lip nt 
genital cone conical; posterior lip with paired 
projections and cuticular inflation of internal 
surface of bursa on eidier side. 

Female. Measurements of 8 specimens, types. 
Total length 3.56-4.81 (4.39); maximum widtii 
0,31-0.48 (0.4)); buccal capsule 0.008-0.010 
(0.009) x .033-0.045 (0039); oesophagus 0.35- 
0.44 (0.39); nerve ring to anterior end 01 6-0.2-3 
(0.20). excretory pore to antenor end 0.21-0.43 
(0.34): deirid to anterior end 0.26-0.37 (0.32), tail 
0.1 6-0. 19 (0.17); vulva to posterior end 0.24-0.28 
(0.26); vagina 0.53-0.99 (0.75), egg 0.055-0.07O 
(0 062) x 0.O3O-O.035 (0 031 ). 

Female tail short, conical; vulva immediately 
anterior to anus; vagina elongate, straight; egg 
ellipsoidal. 



I. BEVERIDGE 




FIGURES 24-36. Cloacina erigone sp. nov., types. 24. Anierior end, lateral view. 25. Cephalic extremity, lateral 
view, dorsal aspect on left-hand side. 26. Cephalic extremity, dorsal view. 27. Cephalic extremity, apical view. 2K. 
Cephalic extremity, transverse optical section through buccal capsule. 29. Submedian cephalic papilla, lateral view. 
30. Preneural oesophageal swelling showing dorsal denticle, dorsal view. 31. Bursa, apical view. 32 Genital cone, 
dorsal view. 33. Spicule tip, lateral view. 34. Gubernaculum, ventral view. 35. Female tail, lateral view. 36. Female 
genital system, lateral view. Scale bars: 24, 31, 35. 36, 0.1 mm; 25-30, 32-34, 0.01 mm. 



CWACINA FROM PAPUA NEW GUINEA 



9 



Remarks 

Ctoucina crigotie sp. nov is characterised by 
having submcdinn cephalic papillae with an 
elongate proximal segment, u shallow, 
unomamented buccal capsule, eight leaf crown 
elements, deirid posterior to the nerve ring, a 
doisal lay with the external branehlets arising 
before the main bifurcation and a straight vagina. 
In these ehaiaclers. C. trigone resembles C. 
cobaUeroi, C. cretheis, C. enyo, C. ips and C. 
sypha.x. It differs from all of these species, 
however, in having bosses lining the anterior part 
of the vagina and a single dorsal denticle. Species 
with a symmetrical buccal capsule, oesophageal 
bosses and a single dorsal denticle arc C. 
au.urahs, C. dis Beveridge. 1998, C. hecuba 
Bevcridgc, 1998. C. io Beveridge, 1998. C lains, 
C lew Beveridge, 1998, C. minor (Davey & 
Wood. I93X) and C. t\ro Beveridge, 1998. 
However, in none of these species is the deirid 
posterior to the nerve ring, and in none do the 
external branehlets of the dorsal ray arise before 
the principal bifurcation. 

This species has, thus far, been found only in 
Do. hagenl. 



Cloacina euryiwme sp, nov. 

(Figs 37-49) 

Types: From stomach of Dendiolugus donanus, 
Tcmbagapura, Irian Jaya, coll. T Flanncry. 
19.V.1994, 23.V.I994. llololype 6, SAM AHC 
31437; allotype ?, SAM AHC 31438. Paratypes. 
4d\99. SAM AHC 31439. Id. 22. SAM AHC 
31440, Id, 1 2, BMNH200L4. 10.5-6, I o\ !?, 
1JSNPC 91135. Slide preparations of spicules, 
apical views of mouth and hursa, SAM AHC 
28381. 

Material examined: From Dettdrolagus 
dorianus: types. From Dendrolagus sctutae: 4 9, 
Swcipini. Sandaun Province. Papua New Guinea, 
coll T Flannery, 15.vi.1991, SAM AHC 31441. 

Description 

Small nematodes; cervical cuticle slightly 
inflated io beyond level of excretory pore, 
transverse cutieular annulalions widely spaced. 
0.030 apart. Submedian cephalic papillae 
prominent, 0.018 long, projecting anteriorly from 
peri-oral cuticle; distal segment ovoid, pointed 
apieally, 0.008 long, shorter than cylindrical 
proximal segment, O.01Q long. Buccal capsule 
shallow, symmetrical in dorsal and lateral views, 
approximately octagonal in transverse section, 



wider than deep, wall without striations. Leaf 
crown elements 8 in number, slightly recurved at 
tips; peri-oral cuticle not inflated into lobes 
attached to each element. Dorsal sector of 
oesophagus bearing opening of dorsal 
oesophageal gland, not projecting into lumen of 
buccal capsule. Oesophagus clavilorm, slender. 
only slightly wider at posterior end, lining 
without sclcrotiscd bosses; Ihree poorly 
developed denticles, one dorsal and two 
subventral, present in lumen of oesophagus 
immediately anterior to nerve ting; prcncural 
swelling of oesophagus small. Nerve ring in mid- 
ocsophageal region; deirids at level of nerve 
ring, excretory pore at level of oesophago- 
intestinal junction. 

Male. Measurements of 7 specimens, types. 
Total length 8.75-10.67 (9.53); maximum width 
0.39-0.48 (0.4S)'. buccal capsule 0.007-O.GlO 
(0.008) x 0.027-0 030 (0.029); oesophagus 0.57- 
0.64 (6.59): nerve ring to anterior end 0.26-0.28 
(0.27); excretory pore to anterior end 54-0.62 
(0.58); deirid to anterior end 0.24-0.32 (028); 
spicules 3.57-3.98 (3 77); gubemaculum 0.015- 
0.020 (0.019) long. 

Dorsal ray elongate, broad at origin, major 
bifurcation occurs at 1/3 length; angle <u 
bifurcation acute; external branehlets arise mid- 
way between major bifurcation and tip, much 
•.homer than internals, directed laterally or postero- 
lateral^, not reaching margin of bursa; internal 
branehlets directed postero-latcrally, almost 
teaching margin of bursa. Extemo-dorsal ray n<« 
reaching matgin of bursa. Gubemaculum 
prominent, wider than long. Spicule tip blunt, 
gently curved; ala terminates abruptly anterior to 
spicule lip. anterior lip of genital cone conical; 
posterior lip with paired projections and cutieular 
inflation of internal surface of bursa on either 
side. 

Female. Measurements of 10 specimens, types. 
Total length 10.19-13.28 (11.50); maximum 
width 0.52-0.75 (0.60); buccal capsule 0.005- 
0.013 (0.008) x 0.030-0.035 (0.031); oesophagus 
0.61-0.73 (0.65); nerve ring to anterior end 0,25- 
0.28 (0.27); excretory pore to anterior end 0.52- 
0.71 (0,60); deirid to anterior end 0.18-0.34 
(0.27); tail 0.22-0.29 (0.25); vulva to posterior 
end 0.31-0 46 (0.39); vagina 1.57-1. 85 (1.75); 
egg 0.08O-O.095 (0.086) x 0.040-0050 (0.046). 

Female tail slender, conical; vulva immediately 
anterior to anus; vagina elongate, straight distally, 
recurrent anterior to vestibule, recurrent section 
twisted in characteristic figure of eight formation; 
egg ellipsoidal. 



10 



I. BEVER1DGE 




FIGURES 37-49. Cloacina euryname sp. nov., types. 37. Anterior end, lateral view. 38. Cephalic extrcmily, lateral 
view, dorsal aspect on right-hand side. 39. Cephalic extremity, ventral view. 40. Submcdian cephalic papilla, lateral 
view. 41. Cephalic extremity, apical view. 42. Cephalic extremity, transverse oplical section through buccal capsule- 
43. Preneural oesophageal swelling showing denticles, lateral view, dorsal aspect on right-hand side. 44. Preneural 
oesophageal swelling showing denticles, ventral view. 45. Bursa, apical view. 46. Spicule tip, lateral view. 47 
Gubernaculum, genital cone and thickenings of spicule sheaths, dorsaJ view. 48. Female tail, lateral view. 49. 
Female genital system, lateral view. Scale bars: 37, 45, 48. 49, 0. 1 mm; 38-44, 46, 47, 0.01 mm. 



CL0AC1NA FROM PAPUA NEW GUINEA 



n 



Remarks 

The presence of three preneural denticles at 
the same level in the oesophagus distinguishes 
C. eurynome from all congeners except C. hero 
Beveridge, 1998, C. hermes Beveridge, 1998 
and C. Iiesiia Beveridge. 1998, all of which are 
parasites of grey kangaroos, Macropus 
fuhginosus (Dcsmarest, 181?) and M- giganteus 
Shaw, 1790 in eastern and southern Australia 
(Beveridge 1998) and C, daveyi Mawson, 1977 
in the wallaroo, M. robustus Gould, 1841. C. 
curynome is distinguished from (he species in 
grey kangaroos in having the deirid at the level 
of the nerve ring rattier than well anterior to it 
and in having eight rather than six leaf crown 
elements. It is distinguished from C daveyi 
which has cephalic papillae with large, globose, 
medially directed distal segments. The denticles 
in C eurynome arc vestigial, a feature found 
only m C bestki. C. eurynome is distinguished 
from all congeners other than C. syplmx, from 
DanopMiilus vanheurni, also from Papua New 
Guinea, in the morphology of the vagina, which 
exhibits a highly characteristic recurrent loop, 
antertor to the vestibule, twisted in a figure of 
eight formation. C. eurynome differs from C, 
syphax in possessing denticles, in having a 
straight rather than a sinuous anterior margin to 
the huccal capsule, and in having the external 
branchlets of the dorsal ray arise after the major 
bifurcation 



Cloacina hecale sp. nov. 
(Figs 50-61) 

Types: From stomach of Dendrolagus 
dcnamir.. Lake Trist, Papua New Guinea, coll. L 
Redmond. 1979. Holotvpc o\ BMNH 
1981.4506; allotype 2, BMNH 1981.4507. 
Paratypes: 49el. 882, BMNH 1981.4508-4535. 
Slide preparations of apical views of bursa and 
mouth. SAM AHC 28383 

Material examined: From Dendrolctgus 
dtinunus. types. 

Description 

Robust nematodes, ceivicul cuticle slightly 
inflated to beyond level of excretory pore; 
transverse cuticuUir annulations widely spaced, 
I). 023 apart, Submedian cephalic papillae 
prominent, 0.017 long, projecting anteriorly from 
peri-oral cuticle, distal segment conical, 0.007 
long, only slightly shorter than cylindrical 
proximal segment. 0.010 long. Buccal capsuie 



shallow, symmetrical in dorsal and lateral views, 
circular in transverse section, wider than deep, 
wall without striations; anterior margin slightly 
undulant Leaf crown elements 8 in numhei, 
slightly recurved at tips; peri-OTal cuticle not 
inflated into lobes attached to each element 
Dorsal sector of oesophagus bearing opening pi 
dorsal oesophageal gland, not projecting into 
lumen of buccal capsule. Oesophagus claviform, 
slender, slightly wider at posterior end; lining 
without sclerotised bosses; denticles absent. Nerve 
ring in anterior oesophageal region; deirid? 
anterior to nerve ring; excretory pore at level of 
oesophago-intestinal ju nction. 

Male. Measurements of 10 specimens, types. 
Total length 9.44- 1 3.86 (11.82); maximum widtH 
50-0.67 (0 60); buccal capsule 0.010-0.013 
(0.01 1) x 0.035-0.040 (0.039). oesophagus 0.76- 
0.95 (0.87); nerve ring to anterior end 0.30-0.32 
(0.31); excretory pore to anterior end (152-0.98 
(0.80); deirid to anterior end 0.17-0.26 (0.23 1 : 
spicules 3.90-5.05 (4.54); gubernaculum 0.040 
long. 

Dorsal ray broad at origin; major bifurcation 
occurs at 1/2 length, external branchlets arise 
immediately after major bifurcation, as long as 
internals, though more robust, directed postero- 
lateral^, nui reaching margin of bursa; internal 
branchlets diiectcd posteriorly almost reaching 
margin of bursa. Externo-dorsal ray not reaching 
margin of bursa. Gubernaculum prominent, wider 
lhan long. Spicule minutely bifid at tip; ala 
terminates abruptly anterior to spicule lip: anteiioi 
lip of genital cone conical; postetior lip with 
paired projections and cuticular inflation of 
internal surface of bursa on eidicr side. 

Female. Measurements of 5 specimens, types. 
Total length 19.6-22.9 (21.3) maximum widdi 
0.64-0.91 (0.81); buccal capsule 0.010 (0.010) x 
0.040 (0.040); oesophagus 0.95-F02 (0.97); 
nerve ring to anterior end 0.25-0.29 (0.27). 
excretory pore to anterior end 0.63-0.86 (0.70); 
deirid to anterior end 0.19-0.29 (0.22). tail 0.13- 
0.30 (0.21); vulva to posterior end 0.18-0 43 
1 1) .28); vagina 2.22-Z57 (2.40); vestibule 0.27; 
sphincter 0.22: mfundibulum 0.17; egg 0.090- 
0.1 10 (0.096) x 0.045-0 050 (0 048). 

Female tail short, conical; vulva immediately 
anterior to anus; vagina elongate, straight, extends 
slightly anterior to vestibule with short recurrent 
section, egg ellipsoidal 

Remarks 

Cloacina hecale sp. nov. Is a robust spccie.% 
characterised by long spicules and an elongate 



1. BEVERIDGE 




FIGURES 50-61. Cloacina hecale sp. nov., types. 50. Anterior end, lateral view. 51. Cephalic extremity, lateral 
view, dorsal aspect on right-hand side. 52. Cephalic extremity, dorsal view. 53. Submedian cephalic papilla, lateral 
view. 54. Cephalic extremity, apical view. 55. Cephalic extremity, transverse optical section through buccal capsule. 
56. Bursa, apical view. 57. Spicule tip, lateral view. 58. Gubernaculum. ventral view. 59. Genital cone, dorsal view. 
60. Female tail, lateral view. 61. Female genital system, lateral view. Scale bars: 50, 56, 60, 61, 0.1 mm; 51-55, 
57-59, 0.01 mm. 



CLOACISA PROM PAPUA NEW GUINEA 



13 



vagina, slightly recurrent at its anterior extremity. 
The length of the vagina relative to the sizes of 
vestibule, sphincter and infundibulum -are given 
in the description in this case to emphasise the 
extreme length of the vagina. The other features 
of the species are unremarkable, with cephalic 
papillae bearing a distal segment with an acute 
tip, 3lmosl equal in length to the proximal 
segment, eight leaf crown elements, a 
symmetrical buccal capsule, a slender, 
unornamcnied oesophagus, Ihe deirid anterior to 
the nerve ring aud the external branchleis of ihe 
dorsal ray arising after the principal bifurcation. 
These features together with the length of ihe 
spicules (:> 3.0 mm) differential C. hecale from 
congeners except C. clymeuc Bevcridge, 1998, 
C aula Johnston & Mawson, 1938. C. liebigi 
Johnston & Mawson, 1 938, C. longispiculata 
Johnston & Mawson. 1939. C. nike Beveridge. 
I99H. C robertsi Johnston & Mawson. 1939, C. 
IWtiUfSfle Mawson, 1975 and C. solymus. C. 
hecale is differentiated from C. clymene and C 
robertsi which have six elements to the leal 
crown and lip-like inflations of the peri-oral 
ctuicle attached to each element of the leaf 
crown. C. curia, C. liebigi, C. longispkulaia and 
C. smalesae all have six leaf crown elements 
rather than eight and the vagina is prominently 
recurrent in these species rather than having a 
short anterior recurrent section- In addition, C, 
liebigi. C, longispiculata and C. smalesae have 
the cervical cuticle inflated so as to form 
'shoulders' in the oesophageal region C, nike 
has submediau cephalic papillae in which the 
distal segment is much shorter than the proximal 
segment and has a sinuous vagina and a longer, 
slender tail in the female. C. solymus has a 
robust subcylindrical oesophagus but the 
submedian cephalic papillae are tiny and the 
anterior margin of the buccal capsule is undulate. 
Therefore. C. hecale is readily distinguishable 
from all known congeners. 



Cloactna hyperea sp. nov. 
(Pigs 62-74) 

Types: From stomach of Dorcopsis hageni. 
Usmo. Madsing, Papua New Guinea, coll. T. 
Rcardon. May, I9K7. Holotype J. SAM AHC 
31442. allotype 9, SAM AHC 3 1443 Paratopes: 
12.5. 82, SAM AHC 31444, \6, 12, BMNH 
2001.4. 10.7-8. Id, 12, USNPC 91136. Slide 
preparations of apical views of mouth and bursa, 
SAM AHC 28382. 



Material examined: Fiom Dorcopsis hagem: 
types. 

Description 

Robust nematodes; cervical cuticle slightly 
inflated to beyond level or excretory pore; 
transverse cuticular simulations widely spaced, 
0.034 apart. Suhmcdian cephalic papillae 
prominent, 0.015 long, projecting anteriorly from 
peri-oral cuticle; distal segment conical. 0.007 
long, only slightly shorter than cylindrical 
proximal segment, 0.008 long. Buccal capsule 
shallow, symmetrically arched anteriorly in lateral 
views; in dorsal views, buccal capsule arches 
anteriorly over dorsal oesophageal gland; in 
ventral views, curves posteriorly; oval and dorso- 
ventrally elongate in transverse section, wider than 
deep, wall with prominent strialions; anterior 
margin smooth, except on dorsal aspect. Leaf 
crown elements 6 in number, slightly recurved at 
tips; peri-oral cuticle slightly inll.Uf.cl into lobes 
attached to each element. Dorsal sector ol 
oesophagus bearing opening of dorsal 
oesophageal gland, projecting prominently into 
lumen of buccal capsule. Oesophagus clavifbrm, 
robust; lining with sclerotised bosses extending to 
level of nerve ring; single dorsal oesophageal 
denticle present in prominent preneural swelling. 
Nerve ring in mid-oesophageal region; deirids at 
level of nerve ring; excretory pore at level of, or 
slightly anterior to, oesophago-iritcslinal junction 

Male. Measurements of 10 specimens, types. 
Total length 4.20-5.21 (4.56); maximum width 
0.26-0.40 (0.34); buccal capsule 0.013-0.020 
(0.016) x 0,055-0.060 (0.056); oesophagus 0.42- 
0.50 (0.47V; nerve ring to anterior end 0.20-0.24 
(0.22); excretory pore to anterior end 0.35-0.51 
(0.43); deirid to anterior end 025-0.34 (0.31); 
spicules 2.25-2 49 (2.37); gubernaculum 0.020- 
0.030 (0.026) long. 

Dorsal ray broad at origin; major bifurcation 
occurs at 1/2 length; angle of bifurcation acute; 
external branchleis arise after major bifurcation, 
as long as internals, directed postero-laterally, net 
reaching margin of bursa; internal branchleis 
directed posteriorly, not reaching margin of bursa. 
Extcmo-dorsal ray not reaching margin of bursa. 
Gubernaculum prominent, slightly wider than 
long. Spicule tip blunt; ala diminishes gradually 
in width towards spicule tip; anterior lip ol genital 
cone conical; posterior lip witfi paired projections 
and cuticular inflation of internal surface of bursa 
on either side. 

Female. Measurements of 6 specimens, types. 
Total length 5 25-6.14 (5 67); maximum width 



M 



I, BEVER1DGE 




FIGURES 62-74. Cloacina hy/ierea sp. nov., types. 62. Anterior end, lateral view. 63. Cephalic extremity, lateral 
view, dorsal aspect on right-hand side. 64. Cephalic extremity, dorsal view. 65. Cephalic extremity, ventral view. 
66. Submedian cephalic papilla, lateral view. 67. Cephalic extremity, apical view. 68. Cephalic extremity, transverse 
optical section through buccal capsule. 69. Preneural swelling of oesophagus, dorsal view, showing denticle. 70. 
Bursa, apical view. 71. Spicule lip, lateral view. 72. Gubernaculum and genital cone, dorsal view. 73. Female tail. 
lateral view. 74. Female genital system, lateral view. Scale bars: 62, 70, 73. 74. 0.1 mm; 63-69, 71.72, 0,01 mm. 



CLUACWA FROM PAPUA NEW GUINEA 



15 



0.32-0.45 (0.39); buccal capsule 0.013-0.018 
(0.016) x 0.O5X-0.060 (0.059j; oesophagus 0.46- 
0,52 t0.49); nerve ring to anterior end 20-0.23 
(0.22); excretory pore Id anterior end 0.35-0.50 
(0.45); deirid to anterior end 0.24-0.33 (0.29); tail 
0. 19-0.26 (0.23); vulva to posterior end 0.36-0.43 
t0-39): vagina 0.96-1.24 (1.07). 

Female tail slender, conical; vulva immediately 
anterior to anus; vagina elongau\ sinuous, extends 
slightly anterior to vestibule vvuh short recurrent 
section; egg not seen. 

Remarki 

The anteriorly arched buccal capsule wall 
distinguishes G hyperea sp. nov. from all 
congeners except C circe Bevcridge, 1999 and C. 
Iaius Beveridge. 1999, both of which occur in the 
quokka, Setonix brachyums (Quoy & Gaimard, 
i 830) in the south-west of Western Australia, and 
from C. nephele sp. nov., described below. C. 
hyperea is distinguished from C. circe in 
possessing selerotised bosses and a dorsal denticle 
in the oesophagus. It is differentiated from C. 
Iaius in having the buccal capsule less 
prominently arched, in having submedian cephal it- 
papillae of a different shape, with both segments 
ol approximately equal length, whereas in C. luius 
the proximal segment is almost twice as long as 
the distal segment. In addition, the spicules are 
2.25-2.49 mm long in C. hyperea compared with 
1.50—1.97 mm in C. Iaius and the vagina is 
commcnsuraiely longer, being 0.96-1.24 mm in 
C, hyperea compared with 0.71-0.92 mm in C. 
taius. In C. hyperea the internal and external 
bianehiets of the dorsal ray are of approximately 
equal length whereas in C. Iaius ihe external 
branchlets are much shorter than the internal 
branchlets C. nephete sp. nov. has lateral lips, 
which are absent in C. hyperaea, and a bulbous 
proximal segment to the cephalic papilla. 



Cloacitia nephele sp. nov. 

(Figs 75-89) 

Tyfi&lt'. from stomach of Darcopsis hageni. 
Usino, Mudang, Papua i\ T ew Guinea, coll. T. 
Rcardoru May, 1987. Hnlotype $, SAM AHC 
31445; allotype V, SAM AHC 31446. Paralypcs; 
3d. 2i\ SAM AHC 31447, Id, BMNH 
2001.4.10.9, \<f, USNPC 91137. Slide 
preparations ol apical views of moulh and bursa, 
SAM AHC 28384. 

Material examined: From Dorcopsis hageni: 
types. 



Description 

Robust nematodes, cervical cuticle prominently 
inflated to beyond level ot excretory pore, 
transverse cuticular annulations widely spaced. 
0.040 apart. Submedian cephalic papillae 
prominent, 0.020 Jong, projecting anteriorly from 
inflated pen-oral cuticle; dist3l segment slender, 
conical, 0.010 long, as long as robust, 
asymmetrical proximal segment, 0.010 long. 
Buccal capsule shallow, symmetrically arched 
anteriorly in lateral views: dorsally, arches 
anteriorly over dorsal oesophageal gland; 
ventrally, curves posteriorly; oval and dorso- 
vent rally elongate in transverse section, wider than 
deep, wall with prominent striations; anterior 
margin smooth. Leaf crown elements 6 in number, 
recurved at tips; peri-oral cuticle inflated into Hp- 
tikc lobes attached to each element; extra dorsal 
and ventral projections of peri-oral cuticle present, 
separate from lateral arcades ol amphids and 
submedian papillae, giving the appearance of lips. 
Dorsal sector of oesophagus bearing opening of 
dorsal oesophageal gland, not projecting into 
lumen of buccal capsule. Oesophagus claviforrn, 
robust: lining without selerotised bosses, denticles 
ahsent in preneural region. Nerve ring in mid- 
oesophageal region; deirids at level of nerve ring; 
excretory pore between nerve ring and oesophago- 
intestinal junction. 

Male. Measurements of 5 specimens, types 
Total length 6.08-7.40 (6.66). maximum width 
0.33-0.37 (0.35); buccal capsule 0.023-O.O30 
(0 026) x 0.090 (0.090); oesophagus 0.69-O W 
(0.71); nerve ring to anterior end 0.28-0.30 
(0.29); excretory pore to anterior end 0.48-0.53 
(0.51), deirid to anterior end 31-0J6 (0.34): 
.spicules 1.96-2.10 (2.03), gubernaculum 035- 
0.040 (0.039) long. 

Dorsal ray bToad at origin, narrowing 
posteriorly, major bifurcation occurs at 1/2 length: 
angle ol bifurcation acute; external branchlets 
arise after major bifurcation, shorter than 
internals, directed postero-laterally. not reaching 
margin ol bursa; internal branchlets directed 
posteriorly, almost reaching margin of bursa. 
Externo-dorsal ray not reaching margin of bursa. 
Gubernaculum prominent, slightly wider than 
long. Spicule tip blunt; ala diminishes gradually 
in width towards spicule lip; anterior lip of genital 
cone conical; posterior lip with paired projections 
and cuticular inflation of Internal sulfate of bursa 
on either side. 

Female. Measurements of 3 specimens, types. 
Total length 5.94-7.87 (7.02): maximum width 
0.34-0.63 (0.50): buccal capsule 0.025-0.030 



16 



I. BEVER1DGE 




FIGURES 75-89. Cloacina ncphele sp. nov., types. 75. Amerior end, lateral view. 76. Cephalic extremity, lateral 
view, dorsal aspect on right-hand side. 77. Cephalic extremity, lateral view, dorsal aspect on right-hand side, 
median optical section showing leaf crown elements and inflation of cephalic collar. 78. Cephalic extremity, dorsal 
view. 79. Cephalic extremity, ventral view. 80. Cephalic extremity, ventral view, median oplical section showing 
leaf crown elements. 81. Submedian cephalic papilla, lateral view. 82. Cephalic extremity, apical view. 83. Cephalic 
extremity, transverse optical section through buccal capsule 84. Bursa, apical view. 85. Genital cone, dorsal view. 
86. Gubcrnaculum, ventral view. 87. Spicule tip, lateral view. 88. Female tail, lateral view. 89. Female genital 
system, lateral view. Scale bars: 75, 84, 88, 89. 0.1 mm; 76-83. 85-87, 0.01 mm. 



CWACINA FROM PAPUA NEW GUINEA 



17 



(0.027) x 0.101) (0)00); oesophagus 0.72-0.79 
(0 76); nerve ring to anterior end 0.29-0.32 
(O.30); excretory pore io anterior end 0.46-0.58 
(0 52). deind to anterior end 0.30-0.31 (0.31); tail 
3 1 -O 39 (0.35); vulva to posterior end 0.43-0.55 
(0 48); vagina 0.51-0.66 (0.59); egg 0.080-0.085 
(0.083) x 0.040-0.045 (0.043). 

Female tail slender, conical, vulva immediately 
anteiioi to anus, vagina elongate, sinuous, extends 
slightly anterior to vestibule with short recurrent 
section; egg ellipsoidal. 

Remark* 

C. nephete sp. uiiv. closely resembles C. circe, 
C. luiun and C. hyperea in having an anteriorly 
arched buccal capsule, li differs from C. laitlf and 
C. hyperea ill lacking Oesophageal bosses and 
denticles, and differs from C. circe in having a 
markedly inflated cervical cuticle, cephalic 
papillae in which the distal segment is not 
oriented medially, and a recurrent vagina. It 
differs from all of these species in having a 
swollen cephalic collar and lip-like inflations of 
the perioral cuticle attached to each leaf crown 
clement. The dorsal and ventral 'lips' are unique 
within the genus. 



< i'weina oneni sp, nov. 
(Figs 90-101) 

Synonyms: Claacina Sp. nov. of Beveridge, 
1998, p 506 (Mai ropus a%i[is). 

Types: From stomach of Mac ropus cujUis, Bula 
Plain. Bcnsbach. Papua New Guinea, coll. I. 
Owen. May. 1998. Holotypc <•?. SAM AHC 
3I44H; allotype 9, SAM AHC 31449. Paratypes: 
51c", 55?, SAM AHC 31450. Id, 1$. BMNH 
1998.9.28.11-12. 

More/ia! examined: From Macrvpus ttgfttSi 
types; Itf, 19, Darl, Bcnsbach. Papua New 
Guinea. SAM AHC 11719. 

Description 

Robust nematodes; cervical cuticle slightly 
inflated to level of excretory pore; transverse 
cuiiculat annotations widely spaced, 0.030 apart. 
Submedian ccphaJic papillae prominent, 0.013 
long, projecting anteriorly from inflated pen-oral 
cuticle; distal segment ovoid, 0.004 long, shorter 
than cylindrical proximal segment. 0.008 long. 
Buccal capsule shallow, symmetrical in lateral and 
dorso-ventral views; octagonal in transverse 
section, wider than deep, wall without prominent 
striaiions; anterior margin smooth. Leaf crown 



elements 8 in number, recurved at lips; peri-oral 
cuticle inflated into lip-like lobes attached to each 
element. Dorsal sector of oesophagus bearing 
opening of dorsal oesophageal gland, nol 
projecting into lumen of buccal capsule. 
Oesophagus claviform, robust; lining withoul 
sclerotised bosses; single dorsal denticle present 
at level of nerve ring; preneural swelling absent. 
Nerve ring in mid-oesophageal region; deirids in 
anterior oesophageal region, excretory port- 
between nerve ring and oesophago-jntestinal 
junclion. 

Mate. Measurements of 10 specimens, types. 
Total lenglh 5.48-7.50 (6.73). maximum width 
0.26-0.43 (0.34); buccal capsule 0.015-0 025 
(0.019) x 0.045-0.070 (0.056); oesophagus 0.53- 
0.65 (0.60); nerve ring to anterior end 0.25-0.32 
(0.27); exeretory pore to anterior end 0.26-0.53 
(0.44); deirid to anterior end 0.10-0.16 (0.J2); 
spicules 2.19-2.67 (2.46)t gubernaculum 0,025- 
0,040 (0.031) long. 

Dorsal ray broad at origin; major bifurcation 
occurs at 1/3 lenglh; angle of bifurcation obtuse; 
external branchlets arise after major bifurcation, 
near extremity of ray, slightly longer and more 
robust than internals, directed posiero-laterally, 
not reaching margin of bursa; internal branchlets 
directed pustero-lalerally, not reaching margin of 
bursa. Extemo-dorsal ray not reaching margin of 
bursa Gubernaculum prominent, slightly wider 
than long Spicule tip blunt, slightly recurved; ala 
diminishes gradually in width towards spicule tip; 
anterior lip of genital cone conical; posterior lip 
with paired projections and cutieular inflation of 
internal surface of bursa on either side. 

Female. Measurements of 10 specimens, types. 
Total length 6.70-9.36 (8 06); maximum width 
0.30-0.61 (0.46); buccal capsule 0.015-0.025 
(0.020) x 045-O.075 (0.057), oesophagus 0.55- 
0.76 (0.64); nerve ring to anterior end 0.24-0.33 
(028); excretory pore to anterior end 0.32-0.61 
(0.48); deirid to anterior end 0.09-0.16 (0.1 1); tail 
0.14-0.21 (0.17); vulva to posterior end 0.27-0.34 
(0.31); vagina 0.45-0.70 (0.58); egg 0.090-0. 100 
(0.093) x 0.040-0.050 (0.048). 

Female tail short, conical; vulva immediately 
anterior to anus; vagina elongate, convoluted, not 
recurrent, egg ellipsoidal. 

Ketiuirks 

The prominent lips and symmetrical buccal 
capsule distinguish C. oweni sp. nov. from all 
congeners except C. urtemis Beveridge, 1998, C 
caenis Beveridge, 1998, C. clymetie, C 
dindymene Beveridge, 1998, C hypsipyle, C 



18 



1. BEVERIDGE 




FIGURES 90-101. Cloacina oweni sp. nov.. types. 90. Anterior end, lateral view. 91. Cephalic extremity, lateral 
view, dorsal aspect on left-hand side. 92. Cephalic extremity, dorsal view. 9.V Cephalic extremity, apical view. 94. 
Cephalic extremity, transverse optical section through huccal capsule. VS. Submedian cephalic papilla, lateral view. 
96. Prencural region of oesophagus, showing denticle, dorsal view. 97. Bursa, apical view. 98. Genital cone, dorsal 
view and gubernaculum. 99. Spicule lip, lateral view. 100. Female tail, lateral view. 101 Female genital system, 
lateral view. Scale bars: 90, 97, 100, 101, 0.1 mm; 91-96, 98, 99, 0.01 mm. 



CLOACiNA FROM PAPUA NbW GUINEA 



19 



linstowi Johnston & Mawson, 1940, C main 
Beveridgc, 1998, C. pana, C robertsL C 
smalesae, C. thelidi.s Johnston & Mawson. 1939 
and C. wutlahiae Johnston & Mawson, 1939. The 
presence of a single dorsal denticle in the 
oesophagus and the lack of oesophageal bosses 
distinguishes it from all of these species except C. 
dindxmene. It differs from C. dindymene in the 
shape of the submedian cephalic papillae which 
have elongated dislal segments in C. dindymate, 
and in the shape, of the dorsal ray. the bifurcations 
of which form an acute angle in C dindymene 
rather than the obtuse angle seen in C. oweni. In 
features of the head, C. oweni is most likely to be 
confused wjth C. rnhertsL found in rock wallabies 
of the genus t'etr<igale Gray, 1837 in eastern 
Australia. C. oweni is readily differentiated, 
however, by the oesophageal denticle and the lack 
of a prominently iccurrcnt vagina as exhibited by 
C. roberni. 

C. oweni is abundant in the stomachs of agile 
wallabies in the Bula Plain region of Papua New 
Guinea, but has not been found in the same host 
species in northern Austtalia in spite ol relatively 
intensive examination of this host in all of die 
northern states (Speare et al. 1983; Beveridgc et 
al. 1998). More surprising is the close 
resemblance of this, species to congeners 
occurring in rock wallabies in Auslralia, a 
relationship for which no simple explanation 

currently exists 



Chacina papuensis sp. nov. 
(Figs 102-1 15) 

Types. From stomach of Macropus a^ili.t, Bula 
Plain, Bensbach, Papua New Guinea, coll. I. 
Owen, May, 1998. Holotype <J, SAM AHC 
31451; allotype 2, SAM AHC 31452. Paratypes. 
5cJ. 62, SAM AHC 31453, lo*. IV, BMNH 
1998.9.28.13-14. Slide pteparations of apical 
view of mouth and bursa, SAM AHC 28386. 

Material examined- From Mairopm agilis. 
types. 

Description 

Small nematodes; cervical cuticle not intl.ued; 
transvci.se cuticular annulations widely spaced, 
0.010 apart Submedian cephalic papillae small, 
conical. 0.006 long, projecting anteriorly from 
peri-oral culicle; distal segment ovoid, 0.002 long. 
shorter than cylindrical proximal segment, 0,004 
long. Buccal capsule shallow, symmetrical in 
lateral and dorso-ventral views; circular in 



transverse section, wider than deep, wall without 
prominent striations; anterior margin smooth. Leaf 
crown elements 8 in number, recurved ar tip*; 
peri-oral cuticle not inflated into lip-like lobe* 
attached lo each element. Dorsal sector of 
oesophagus bearing opening of dorsal 
oesophageal gland in prominent, bilobed, tooth- 
like structure projecting into lumen of buccal 
capsule, each ventral sector with 2 triangular 
sclerotised projections. Oesophagus claviform, 
slender; lining without sclerotised bosses; single 
dorsal denticle present immediately anterior to 
nerve ring, preneural swelling present Nerve ring 
in mid-oesophageal region; dcirids in anterior 
oesophageal region: excretory pore between nerve 
ring and ocsophago-intestinal junction. 

Male. Measurements of 5 specimens, types. 
Total length 3 18-5.09 (4.57); maximum width 
0.18-0.32 (.0.23); buccal capsule 0.007 -O.OiO 
(0.008) x 0.02O-O.026 (0.021); oesophagus 0,31- 
0.41 (0,36); nerve ring to anterior end 16-0-17 
(0.17); excretory pore to anterior end 0. 20-0, 34 
(0.28); deirid to anterior end 0,08-0.11 (0.10); 
spicules 1.84-2.26 (2.05); gubernaculum 0.025- 
0.030 (0.029) long. 

Dorsal ray broad at origin; major bifurcation 
occurs at 1/2 length, angle of bifurcation acute:, 
external branchlets arise aftei major bifurcation., 
shorter and more robust than internals, directed 
postcro-laterally. not reaching margin at burva; 
internal branchlets directed posteriorly, noL 
reaching margin of bursa. Extcrno-dorsal ray not 
teaching margin of bursa. Gubernaculum 
prominent, slightly wider than long. Spicule tip 
bifid; ala diminishes gradually in width towards 
spicule lip; anterior lip of genital cone conical; 
posterior lip with paired projections and cuticular 
inflation of internal surface of bursa on cither 
side. 

Female. Measurements of 5 specimens, types. 
Total length 5.14-6.05 (5.57); maximum width 
0.25-0.36 (0.32); buccal capsule 0.005-0 008 
(0.007) x 0.020-0.025 (0.023); oesophagus 0.35- 
0.39 (0.38); nerve ring to anterior end 0.17-0.1* 
(0.17); excretory pore lo anterior end 0.23-0.38 
(0.33); dejrid to anterior end 0.07-001 1 (0,09); 
tail 0.14-0.22 (0.19); vulva to posterior end 0.30- 
044 (0.39); vagina 0.47-0.63 (0.53); egg 0.O7O- 
0.085 (0.080) x 0.040-0.050 (0.046). 

Female tail short, conical; vulva immediately 
anterior to anus; vagina elongate, convoluted, noi 
recurrcnl; egg ellipsoidal. 

Remarks 
The simple, symmetrical buccal capsule, lack of 



20 



1. BEVERIDGE 



102 




FIGURES I02-I15. Cloacina papuensis sp. nov., types. 102 Anterior end. lateral view. 103. Cephalic extremity, 
lateral view, dorsal aspect on left-hand side. 104. Cephalic extremity, dorsal view. 1(15. Submedian cephalic papilla, 
lateial view. 106. Cephalic extremity, apical view. 107. Cephalic extremity, transverse optical section through 
buccal capsule. 108. Preneural region of oesophagus, showing denticle, lateral view, dorsal aspect on left-hand side. 
109. Preneural region of oesophagus, showing denticle, dorsal view. 110. Spicule tips, ventral view. Ill, Bursa, 
ventral view. 112. Bursa, lateral view. 113. Female tail, lateral view. 114. Female genital system, lateial view. 115. 
Genital cone, dorsal view. Scale bars: 1 13. 1 14, 0,1 mm; 102-104, 106-1 12, 115 0.01 mm; 105, 0.003 mm. 



CLOACINA FROM PAPUA NEW GUINEA 



21 



prominent lips and unoniamcntcd oesophagus 
with :i single dorsal denticle differentiate C, 
popuensis sp. nov, from all congeners except C. 
cotnuta, C dure. C. longispiculata and C. scinm. 
C. pcijiuensis is differentiated from C. sciron 
primarily in having the deirid anterior lo, rather 
than at the level of, the nerve ting. In C. scinm, 
the anterior margin ol Ihe buccal capsule arches 
anteriorly and is sinuous. C. papuensis differs 
from C. longispiculata in lacking the prominent 
shoulder-like inflations of the cervical cuticle, in 
having Ihe excretory pore at Ihe level of the 
ocsophago-intcstinal junction rather than well 
posterior to it as in C. longispiculata, and in 
having eigbl Km! crown elements rather than the 
six present in C. longispiculata. It differs from C, 
dircc iii lacking the prominent cervical inflation 
of the cuticle, in Ihe shape of (he submedian 
papillae which have an extended distal segment in 
C. dirce. in spicule lengths (1 ,84-2.26 (2.05) mm 
long in C, papuensis, 3.48-3.95 (3.70) mm long 
in C dine), and in the shape of the vagina which 
is longer and more convoluted in C. dirce. 
Therefore, C. papuensis most closely resembles 
C. cornula, also a parasite of Mocropus agilis, in 
having a prominent dorsal oesophageal tooth. It 
differs from C cornula in lacking a cervical 
cuticulai inflation, in the shape of the submedian 
papillae which have elongate distal segments in 
C. cotnutu, in the shape of the dorsal ray which In 
C. cormtia terminates in very short subequal 
internal and external branchlcls, in spicule length 
(1.84-226 (2.05) in C, papuensis, 1.38-1.62 
(151) mm in C. cornuta), and in the length and 
shape of the female tail which is very short and 
prominently swollen in C. cornula 



Cloacina polymeLi sp. nov. 
(Figs 116- 128) 

Types: From stomach of Oorcopsis hageni. 
Usino, Madang, Papua New Guinea, coll T. 
Reardon. May, 1987. Holotypc 6, SAM AHC 
31454: allotype 9. SAM AHC 31455. Paratypes: 
4J. 8 9, SAM AHC 31456, Id, I ?,. BMNH 
2001.4,10,10-11, IS, 19, USNPC 91138. Slide 
preparations of apical views of mouth and bursa. 
SAM AHC 28387. 

Material examined: From Oorcopsis hageni: 
types. 

Oc v< upturn 

Robust nematodes: anterior extremity deviated 
dorsjilly, cervical cuticle inflated to level of 



excretory pore; transverse cutkular annulations 
widely spaced, 0.030-0.062 apart. Submedian 
cephalic papillae prominent, 0.016 lony, 
projecting anteriorly from inflated pen-oral 
cuticle; distal segment acute, conical, 0.010 long, 
longer than cylindrical proximal segment. 0.006 
Inng. Cephalic collar prominently inflated on 
dorsal and ventral aspects, anterior margin ol 
collar rugose; amphids conical, projecting above 
level of cephalic collar. Buccal capsule very 
shallow, symmetrical, anterior margin highly 
sinuous; due to dorsal deviation of anterior 
extremity, buccal capsule appears to arch dorsally 
in ventral views; oval and dorso-ventrally elongale 
in transverse section, wider than deep, wall finely 
striated. Leaf crown elements 8 in number, 
recurved at tips; peri-oral cuticle inflated into Jip- 
like lobes attached lo each element. Dorsal sector 
of oesophagus bearing opening of dorsal 
oesophageal gland, nol projecting into lumen of 
buccal capsule. Oesophagus claviform, robust, 
anterior section of greater diameter than region 
posterior to nerve ring; lining without sclerolised 
bosses; denticles absent in preneural region. Nerve 
ring posterior to mid point of oesophagus, deirids 
slightly anterior to level of nerve ring; excretory 
pore between nerve ring and oesophago-inlcsfinal 
junction. 

Male. Measurements of 9 specimens, types. 
Total length 5.86-6.03 (5.93 1: maximum widtn 
0.29-0.46 (0.39); buccal capsule 0.020-0. 030 
(0.O25) x 0.115-0.150 (0.130); oesophagus 0.66- 
0,79 (0.72); nerve ring to anterior end 044-0.53 
(0.48); excretory pore to anterior end 0.55-0.70 
(0.65), deind to anterior end 0.36-0.45 (0.3y); 
spicules 2.61-2.71 (2.65); gubemaculum 0.035- 
0.050 (0.045) long. 

DoTsal ray broad at origin; major bifurcation 
occurs at 1/2 length; angle of bifurcation acute: 
external branchlets arise after major bifurcation, 
much shorter than internals, directed postero- 
laterally, not reaching margin of bursa; internal 
hranchlets elongate, directed posteriorly, almost 
reaching margin of bursa Extemo-dorsal ray not 
reaching margin of bursa. Gubemaculum 
prominent, subcordate. Spicule tip blunt, slightly 
recurved; ala diminishes gradually in width 
towards spicule tip; anterior lip of genital cone 
conical; posterior lip wilh paired projections and 
cuiicular inflation of internal surface of bursa on 
either side. 

Female. Mcasuieiuciiis ol 10 specimens, types. 
Total length 5.45-8.22 (7.14); maximum width 
0.33-0.59 (0.46); buccal capsule 0.015-0.030 
(0.025) x 0.120-0 160 (0.150); oesophagus 070- 



22 



I. BEVE.RIDGB 




124 



FIGURES 1 16-128. Cloacina polymela sp. nov., types. 116. Anterior end, lateral view, showing dorsal deviation of 
head. 117. Cephalic extremity, lateral view, dorsal aspect on left-hand side. 118. Cephalic extremity. Ventral view. 
119 Cephalic extremity, dorsal view 120. Cephalic extremity, apical view. 121. Cephalic extremity, transverse 
optical section through buccal capsule. 122. Submedian cephalic papilla, lateral view. 123. Amphid, lateral view. 
124. Bursa, ventral view. 125. Spicule tip, ventral view. 126. Gubernaculum and genital cone, dorsal view. 127. 
Female tail, lateral view. 128. Female genital system, lateral view. Scale bars: 116, 124, 127, 128, 0.1 mm; 117- 
123, 125,126,0.01mm. 



CLOAC1NA FROM PAPUA NEW GUINEA 



23 



0.94 (0.S2); nerve ring to anterior end 0.45-0 61 
(0 54); excretory pore to anterior end 0.55-0.85 
(0.75)-, deirid lo anterior end 0.31-0.50 (0.42); tail 
0.2 1-0.30 (0.24); vulva to posterior end 0.33-0.50 
(0 3K); YagiM 0.49-0.68 (060); egg 0.080-0.085 
(0.082) x 0.040-0 045 (0.042). 

Female tail slender, conical; vulva immediately 
anterior to anus; vagina elongate, .sinuous, extends 
slightly anterior to vestibule with short recurrent 
section; egg ellipsoidal. 

Remark* 

Cluucina polymela sp. nov. is a highly 
distinctive species which differs from all 
congeners in having the anterior extremity 
deviated dorsally. This feature appears in every 
specimen and therefore is not likely to be a 
riX&tiofl artefact In addition, it differs from 
congeners except C. dryope Beveridgc, 1998 and 
I 'uppho in having an oesophagus in which the 
prencwral region is of greater diameter than the 
posterior part. C. polymela is distinguished from 
C. dryope in having eight rather than six leaf 
crown elements, in having a mors posterior deind, 
in having the excretory pore anterior to the 
ocsophago-intestinal junction rather than posterior 
to it. and in the shape of the buccal capsule which 
is extremely shallow in C. dryope and in which 
the anterior margin has only slight saliences rather 
tliaii prominent undulations. Gi polymela and C. 
sappho are the only iwo members of the genus in 
Which the arnphids form acutely pointed conical 
projections above the cephalic collar. In all other 
species, the arnphids are dome-shaped and do not 
project obviously beyond the coUar. C. polymela 
also resembles C sappho in the shape of the 
iloTsal ray of the bursa and in possessing eight 
leaf crown elements. However, it differs in having 
prominent inflations of the cephalic collar on the 
dorsal and ventral surface, a feature which 
resembles the lip-like projections present in C. 
nephele. C. polymela also differs from C. sappho 
in spicule length (2.61-2 71 (2.65) mm in C. 
polymela; 1.30—1.50 (1.38) mm in C. sappho) and 
in the shape of the vagina which is recunent in C. 
polymela but short and straight in C. sappho. 



( Umcina pmtiihra sp nov. 
(fi^s 129 MS) 

Types- From stomach of Dorcopsis hageni. 
Dsino, Madang, Papua New Guinea, coll- T 
Rcardon. May. 1987. Holotype d\ SAM AHC 
31457. Pararypes; 2c?. SAM AHC 31458, 1 cT, 



BMNH 2001.4 10.12; 16. USNPC 91139. Slide 
preparations of apical views of mouth and bursa, 
SAM AHC 28388. 
Material examined: From Dorcopsis hagem: 

types. 

Description 

Robust nematodes; cervical cuticle slightly 
inflated to beyond level of excretory pore; 
transverse culicular annulations widely spaced. 
0.015 apart. Submcdian cephalic papillae 
prominent, 0.019 long, projecting anteriorly from 
peri -oral cuticle, distal segment ovoid, 0.008 long, 
deviated medially, slightly shorter than 
asymmetrical, cylindrical proximal segment, 0.01 1 
long; subniedian papillae situated anteriorly on 
cephalic collar; arnphids situated posteriorly, 
below anterior margin of buccal capsule in lateral 
views. Buccal capsule shallow, symmetrical in 
lateral and dorso-vcntral views, wall with faint 
striations; anterior margin undulate, with anterior 
projections associated with each leaf crown 
element. Leaf crown elements 8 in number, net 
recurved at tips; peri-oral euliele inflated into 
lobes attached to each element Dorsal sector of 
oesophagus bearing opening of dorsal 
oesophageal gland, not projecting into lumen of 
buccal capsule. Oesophagus subcylindrical, almost 
claviform at base, robust; lining without 
sclerotised bosses or denticles; preneural swelling 
of oesophagus absent Nerve ring in mid- 
oesophageal region; deirids anterior to nerve nng; 
excretory pore at level of, or slightly anterior to, 
ocsophago-intestinal junction. 

Male. Measurements of 6 specimens. type.*. 
Total length 3.71-5.34 (4.86); maximum width 
0.32-0 38 (0.36); buccal capsule 0.015-0.018 
lO .016) x 0.050-0.055 (0.054); oesophagus 0.66- 
0.78 (0.71); nerve ring to anterior end 0.34-0.38 
(0.36); excretory pore to anterior end 43-0.49 
(0.46); deirid to anterior end 0.27-0.32 (0.30); 
spicules 1.30-1.58 (.1.43); gubernaculum 0.030- 
0.040 (0.033) long. 

Dorsal ray broad at origin, major bifurcation 
occurs at 1/2 length; angle of bifurcation acute; 
external branchlets arise immediately altet majiu 
bifurcation, much shorter than internals, directed 
postero-laterally, not reaching margin of bursa; 
internal branchlets directed posteriorly, not 
reaching margin of bursa. Externo-dorsal ray not 
reaching margin of bursa. Gubernaculum 
prominent, subcordate, slightly wider than long. 
Spicule tip blunt; ala diminishes gradually m 
width Ihen terminates abruptly anterior to spicule 
tip; anterior lip of genital cone conical; posterior 



24 



1. BEVER1DGE 




FIGURES 129-138. Cloacina praxithea sp. nov., lypes. 129. Anterior end. lateral view. 130. Cephalic extremity, 
lateral view, dorsal aspect on right-hand side. 131. Cephalic extremity, dorsal view; arrows indicate amphids. 132. 
Cephalic extremity, apical view; arrows indicate amphids. 133. Cephalic extremity, transverse optical section 
through buccal capsule. 134. Submedian cephalic papdla, lateral view. 135. Gubernaeulum, ventral view, 136. 
Bursa, apical view. 137. Genital cone, dorsal view. 138. Spicule tip, lateral view. Scale bars: 129, 136, 0.1 mm; 
130-135, 137, 138,0.01mm. 



CLOACINA FROM PAPUA NEW GUINEA 



25 



lip with paired projections and cuticular inflation 
of internal surface of bursa on eilher side. 
f-'eimde. Not seen 

Remarks 

Clouctna praxilhea sp. nov. is distinguished 
from all congeners by the position of the amphids 
on the cephalic collar. In other species the 
amphids are a the same level as the submedian 
papillae, anterior to the level of the buccal 
capsule, while in C. praxitliea the amphids, in 
lateral view, are below the level of the anterior 
margin of the buccal capsule (Fig. 130). C. 
parxiliiea is characterised by a symmetrical, lobed 
buccal capsule without bosses or denticles in the 
oesophagus. As such, it has similarities with the 
Australian species C. at minis, C. dryope, C. hebe, 
C. hyi'sipvle, C. linxtowi. C. main. C. thetbdlS and 
C weulflbkte, all of which differ from it m having 
six rather than eight leaf crown elements. Among 
the species known from New Guinea, the features 
of the buccal capsule, as well as the presence of 
eight leaf crown elements, indicate affinities with 
C. svphax. C. xolon, C. xappho and C. solymus. C. 
solymux has tiny cephalic papillae, in striking 
contrast to the prominent papillae of C. pruxtthea, 
while C iapp/w and C. pclytnela have the 
amphids on or forming projections beyond I he 
level of the cephalic collar; C uilnn has a greatly 
inflated cervical cuticle and its spicules are 2 60- 
2.95 (2 81) mm in length compared with 1.93- 
2.05 (1.99) mm in C. praxUhea; C. syphax differs 
in having die lateral branchlets of the dorsal ray 
arise anterior to the principal bifurcation. 
Therefore., C. pruxiihea is readily distinguishable 
from all congeners even in the absence Of females, 
which arc cuncntlv unknown. 



Cloacina procris sp, uov. 
(Figs 139-153) 

Types: From stomach of Darcapsh hugrni 
i J urn, Madang, Papua New Guinea, cnll. T. 
Reardon. May. I'>87. l-iolutype o\ SAM AHC 
31459; allotype 2, SAM AHC .3 1460. Paratypes: 
1<5. 22. SAM AHC 31461 Slide preparations of 
apical -views of nioulh and bursa. SAM AHC 
28389. 

Mulehul examined: From Dnrcopsis hageni: 
types. 

Description 

Robust nematodes; ceivical cuticle slightly 
inflated to level of excretory pore, transverse 



euticular annulations widely spaced, 0.020 apart 
Submedian cephalic papillae small. 0.0035 long, 
projecting anteriorly from peri-oral cuticle; distal 
segment ovoid, 0.0005 long, much shorter than 
cylindrical proximal segment, 0.003 long. Buccal 
cHpsulc shallow, asymmetrical in lateral views, 
deeper on ventral side, slight anterior arching of 
capsule; dorsally. capsule arches anteriorly over 
dorsal oesophageal tooth; vcntrally, curves 
posteriorly; oval and dorso- vcntrally elongate in 
transverse section, wider than deep, wall with 
faint striations; anterior margin smooth. Leaf 
crown elements 8 in number, not recurved at tips; 
peri-oral cuticle very slightly inflated into lobes 
attached to each clement. Dorsal sector of 
oesophagus bearing opening of dorsal 
oesophageal gland, projecting prominently into 
lumen of buccal capsule; each sector of 
oesophagus bears single triangular tooth-like 
structure projecting into lumen. Oesophagus 
daviform, robust; lining with sclcrotiscd bosses 
extending to level of nerve ring; single dorsal 
oesophageal denticle present in prcneural region, 
preneural swelling of oesophagus absent. Nerve 
ring in mid-oesophageal region; deirids posterior 
to nerve ring; excretory pore at level of, or slightly 
anterior to, oesophago-intestinal junction. 

Male. Measurements of 3 specimens, types 
Folal length 3,25-4.52 (3.89); maximum width 
0.17-0.32 (0.26), buccal capsule 0.015-0.01 K 
(0.017) x 0.050-0.060 (0.054); oesophagus 0.40- 
0.41 (0.40): nerve ring to anterior end 0.20-0.21 
(0.21); excretory pore to anterior end 0.35-0.40 
(0.38); deirid to anterior end 0.27-0.32 (0.30); 
spicules 1.30-1 58 (1.43); gubernaculum 0.030 
(0.030) long 

Dorsal lobe of bursa elongate; dorsal ray broad 
at ongin; major bifurcation occurs at 1/2 length; 
angle of bifurcation acute; exLcrnal branchleis 
arise at level of major bifurcation, much shortei 
than internals, directed almost laterally, not 
reaching margin of bursa; internal branchleis 
directed posteriorly, reaching margin of bursa 
Extemo-dorsal ray not reaching margin of bursa. 
Gubernaculum prominent, slightly wider than 
long. Spicule tip blunt; ala diminishes gradually 
in width (hen terminates abruptly anterior to 
spicule lip; anterior lip of genital cone conical; 
posterior hp with paired projections and culiculai 
inflations of internal surface of bursa on cither 
side. 

Female. Measurements of 3 specimens, types 
Total length 4.26-5.15 (.4.64); maximum width 
0.37-0.47 (0.42); buccal capsule 0.018-0.020 
(0.019) x 0.060-0.065 (0.062); oesophagus 



]. BEVERJDGE 




149 



FIGURES 1 39-153. Cloacina pmcris sp. nov., types. 139. Anterior end, lateral view. 140. Cephalic extremity, 
lateral view, dorsal aspect on left-hand side. 141. Submedian cephalic papilla, lateral view. 142. Cephalic extremity, 
dorsal view. 143. Cephalic extremity, ventral view. 144. Cephalic extremity, apical view. 145. Cephalic extremity, 
transverse optical section through buccal capsule. 146. Transverse optical section through anterior extremity of 
oesophagus showing bosses and three tooth-like projections in each of the oesophageal sectors. 147 Preneural 
region ol oesophagus, dorsal view, showing denticle. 148 Gubernaculum. ventral view. 149. Genital cone, dorsal 
view. 150 Bursa, apical view. 151. Spicule tip, lateral view. 152. Female tail, lateral view. 153. Female genital 
-system, lateral view. Scale bars: 139, 152, 153, 0.1 mm; 140, 142-151, 0.01 mm; 141, 2.5 urn. 



CLOACINA PRQM PAPUA NEW GUINEA 



17 



0.48-0 50 (0.49); nerve ring to anterior end 
0.21-0.22 (0.22); excretory pore to anterior end 
0.36-0.40 (0.3R); deirid to anterior end 0.29- 
0.33 (0 31); tail 0.19-0.27 (0.23); vulva to 
posterior end 0,35-0.47 (0.40); vagina 0.52-0.60 
(0.53); egg 0.050-O.060 (0.055) x 0.030-0.035 
(0.032). 

Femaie tail slender, conical; vulva immediately 
anterior to anus; vagina elongate, straight, tutus 
;ibrupily at entry to vestibule; egg ellipsoidal. 

Remarks 

Although described from only a small scries 
of specimens. C, procris sp, nov. is quite 
distinctive morphologically It is characterised 
by very small submedian cephalic papillae, a 
slightly asymmetrical, arched buccal capsule, 
eight leaf crown elements, bosses lining the 
anterior region of the oesophagus, a single, 
small oesophageal denticle, deirid posterior to 
Ihe nerve ring, an elongate dorsal ray with the 
external branchlets arising immediately after the 
principal bifurcation, and a straight vagina. 
These features distinguish the species from all 
congeners. 

The asymmetrical buccal capsule, oesophageal 
bosses and dorsal denticle with the posterior 
position of the deirid distinguish the species from 
all others except C cos Beveridge, 1998, C. 
paptllata Beveridge, 1979 and C. stempe In C. 
60S and C. papillata there are only six rather than 
eight leaf crown elements and (he vagina is 
prominently recurrent. In C. sterope, the buccal 
capsule exhibits a much greater degree of 
asymmetry, the submedian cephalic papillae are 
[fttgcf and more prominent and the spicules arc 
longer (1.67-2.07 (1-96) mm in C. srerope, 1.30- 
1.58 (1.43) mm in C. procris). 



Cloacina stetope Beveridge & Spcare, 1999 

Material examined. 3 c', 4 9, from stomach of 
Den-up six hageni, Ustno, Madang, Papua New 
Guinea, cull. T. Reardon, May, 1987, SAM AHC 
31461 

Remark* 

Cloacina sterope was described by Beveridge 
and Speure {1999) fnim Dorcopsulus vanfiearni 
horn Ihe Chimbu Province of New Guinea. 
Morphological features as well as measurements 
of the new specimens from D. hageni agree well 
with Ihe original description. D. hageni 'represents 
a new host record. 



Cloacina syphax Beveridge & Spcarc, 1999 

Material examined: I7tJ, 9$. from stomach of 
Dorcapsis hageni. Usino. Madang. Papua New 
Guinea, coll. T. Reardon. May. 1987, SAM AHC 
31469;BMNH200I4.I0.13. 

Remarks 

Cloacina syphax was described from 
Dorcopsulus vanheurni by Beveridge and Spearc 
(1999). The current collection represents a new 
host record. 



Cloacina theope sp. nov. 
(Figs 154-166) 

Types: From stomach of Dendrolagus matschici 
Forstcr & Rothschild, 1907, Huon Peninsula. 
Morobe Province, Papua New Guinea, coll. J. 
Mayer, July 1999. Hoiotype o, USNPC 91140. 
allotype 9, USNPC 91141. Paratypes: 2c?, 49, 
USNPC 91 142; I o*. on slides, SAM AHC 28390 

Material examined: From Dendrolagm, 
matschiei: Iypcs. From Dendrolagus dnrianus: 
lOd, 5 9, Lake Trist, Papua New Guinea, coll. ), 
Redmond, 1979, BMNH 1981. 4486-4505. 

Description 

Robust nematodes; cervical cuticle sligluly 
inflated; transverse euticular annulations widely 
spaced, 0.021-0.044 apart. Submedian cephalic 
papillae prominent, 0.017 long, projecting 
anteriorly from inflated peri-oral cuticle; distal 
segment ovoid, 0.008 long, directed slightly 
medially, slightly shorter than robust, 
asymmetrical proximal segment, 0009 long 
Buccal capsule shallow, symmetrical in lateral and 
dorso- ventral views; circular in transverse section, 
wall without prominent striatums, anterior margin 
smooth. Leaf crown elements 8 in number 
recurved at tips, peri-oral cuticle not inflated mio 
lip-like lobes attached to each element. Dorsal 
sector of oesophagus bearing opening of dorsal 
oesophageal gland, not projecting into lumen of 
buccal capsule. Oesophagus clavilorm, slender: 
lining without sclcrorised bosses; denticles present 
in preneural region, dorsal denticle prominent; 
two subventral thickenings of oesophageal lining 
giving appearance of denticles, at same level as 
dorsal denticle. Nerve ring in mid-oesophageal 
region, deinds at level of nerve ring; excretory 
pore at level of oesophago-mtestinaljunction. 

Male. Measurements of 4 specimens, types. 
Total length 4.68-5.04 (4.88); maximum width 



28 



I. BEVERIDGE 




FIGURES 154-166. Cloacina iheope sp. nov., types. 154. Anterior end, lateral view. 155. Cephalic extremity, 
lateral view, dorsal aspect on left-hand side. 156. Cephalic extremity, dorsal view. 157. Submedian cephalic papilla, 
lateral view. 158. Cephalic extremity, apical view. 159. Cephalic extremity, transverse optical section through 
buccal capsule. 160. Preneural region of oesophagus, lateral view, dorsal aspect on left-hand side, showing 
denticles. 161 Preneural region of oesophagus, dorsal view, showing denticles. 162. Bursa, apical view. 163. 
Gubernaculum. genital cone and spicule sheaths, dorsal view. 164. Spicule tip. lateral view. 165. Female tail, lateral 
view. 166. Female genital system, lateral view. Scale bars: 154, 162, 165, 166, 0.1 mm; 155-161, 163-164, 
0.01 mm. 



CWAC1NA FROM PAPUA NEW GUINEA 



29 



0.30-0.32 (0.31): buccal capsule 0.010 (0.010) x 
0.030-0.033 (0.031), oesophagus 0.51-0,60 
(0.55); nerve ring lo -anterior end 0.18-0.20 
(0.19); excretory pore lo anterior end 0.31-0.4] 
(0.35); detrid to anterior end 0.13-0.23 (0.18); 
spicules 2.29-2.63 (2.52); gubcmaculuni 0.015- 
020 (0.01 R) long. 

Dorsal ray broad at origin; major bifurcation 
occurs at 1/2 length; angle of bifurcation obtuse; 
external branchlets; arise immediately after major 
bifurcation, shorter than internals, directed 
laterally, not reaching margin of bursa; internal 
branchlets directed posiero-laicrally, almost 
reaching margin ol bursa. Exlerno-dorsal ray not 
teaching margin of bursa Gubernaculutti 
prominent, slightly wider than long. Spicule tip 
minutely bifid; ala diminishes gradually in width 
towards spicule tip. anterior lip of genital cone 
Conical; posterior lip with paired projections and 
cuticular inflations of internal surface of bursa on 
either side. 

Female. Measurements of 5 specimens, types. 
Totnl length 6.09-6 71 (6.29); maximum width 
0.39-0.43 (0.41K buccal capsule 0.008-0.010 
(0.009) x 0.030-0.035 (0.032); oesophagus 0.52- 
64 (0.59); nerve ring to anterior end 25 (0.25); 
excretory pore lo anterior end 30-0.51 (0.43); 
detrid lo anterior end O.U-0.23 (0.20); tail 0.28- 
0.36 (0.33); vulva to posterior end 0.49-0.68 
(0.59); vagina 1.03-1.33 (1.19); egg 0.085-0.095 
(0.089) x 0.045-0.050 (0 048). 

Female tail slender, conical; vulva immediately 
anterior lo anus; vagina elongate, sinuous, extends 
slightly antenur to vestibule with short recurrent 
section; egg ellipsoidal. 

Remarks 

Cloacina theope sp. nov. differs from 
Congeners in having three oesophageal denticles 
in the prcncural region of the oesophagus which 
are at the same level, but are of dissimilar sizes. 
The dorsal denticle is well developed but the 
subveniral denticles are mere thickenings of the 
oesophageal lining and may not warrant the 
designation as denticles. In all other species with 
(luce denticles at the same level in the 
oesophagus. G (Jbveyi>, C. eurynnmc, C. hera. C. 
hermes, C hfftfO, the denticles are equally 
developed, although in C heslia and C. eurynomc 
all three denticles may be weakly sclerotised. C. 
theope differs from C. daveyi. which has 
suhmedian cephalic papillae in which the distal 
segment is large and directed medially, and from 
all ihe remaining species, which have the deirid in 
ill' anterior oesophageal position and six rather 



than eight leaf crown elements. It is most similar 
10 C. eunnome from which it differs principally 
in having shorter spicules (3.57-3.98 (3.77) mm 
in C. ettrynome, 2.29-2.63 (2.52) mm in C. 
theope). 

If the subveniral denticles are excluded from 
consideration, then the unornamented anterior 
oesophagus, a symmetrical buccal capsule with » 
smooth anterior margin, and the presence of a 
dorsal denticle indicates similarities with C 
cnpuUa, C. dindymene, C dirce, C. ernabella, C. 
longispicultita and C. scirrm. C. ernabella differ* 
in the shape of the cephalic papillae (globose, 
medially directed distal segment in C. ernahrllal 
while all species except C xciron differ in having 
the deirid in the anterior oesophageal region raiher 
than at the level of the nerve ring. C, theopt 
differs from C. sciron. which has an anteriorly 
arched buccal capsule, longer spicules (2.83-3.30 
(3.00) mm in C. sciron, 2.29-2.63 (2.52) mm in 
C. theope), a longer dorsal ray and a vagina which 
is not recurrent. In C. theope the recurrent section 
of the vagina is twisted around the ascending arm 
in a manner similar to that found in C, syphos and 
C eurynome, C theope differs from C sypha.x in 
the shape of the buccal capsule, which has an 
undulating anterior margin in C. zyphax but is 
straight in C. theope. Therefore, C. theope iv 
clearly distinguishable from all congeners. 



Discussion 

The new records and new species reported in 
this paper suggest that a diverse array of species 
of Cloacina exists in macropodid marsupials in 
Papua New Guinea. Conclusions need to be 
guarded as the current collections have been 
obtained from a small number of individuals of 
each host species, in some instances from 3 single 
animal. As yet. there have been no comprehensive 
surveys of New Guinean macropodids for 
parasites, and several species of Thytogale. 
Dorcopsis, Dnrcopsulus and Uendrolagus have 
apparently yet to be examined for helminths 
(Spratt et al. 1991). In addition, Beveridge (1998) 
and Beveridge and Speare (1999) have listed 
museum records of additional undescribed species 
from some of ihese hosts which cannot currently 
he named due to lack of adequate material. 

In spile of Ihe relatively rudimentary knowledge 
of ihe New Guinean parasiie fauna, some 
preliminary comparisons can be made wilh species 
present in Australia. In terms of morphological 
characters, the species of Cloacina present in New 



30 



I. BEVERIDGE 



Guinea are virtually as diverse as those present in 
Australia. Of the various morphological features of 
the genus, only the spiral twisting of the proximal, 
recurrent region of the vagina (C. syphax, C. 
eurynome, C. theope) and the presence of amphids 
on conical projections (C. praxithea, C. sappho) 
appear to be restricted to species from New Guinea. 
Two other apparently autapomorphic characters, 
using Arundelia dissimilis (Johnston & Mawson, 
1939) as an outgroup, are the posterior position of 
the amphids in C. praxithea and the dorsal 
deviation of the head in C. polymela. A. dissimilis 
is the sole member of a closely related genus within 
the tribe Cloacininea (see Beveridge 1987) and is 
therefore considered to be an appropriate outgroup 
for polarising morphological characters. Characters 
such as the origin of the external branchlets of the 
dorsal ray anterior to the major bifurcation are more 
common in species from New Guinea, but are also 
found in species from northern Australia. No 
obvious patterns are detectable in the distribution 
of other characters. While only tentative 
conclusions can be drawn in the absence of a 
formal phylogenetic analysis, there do not appear 
to be any major distinctions which can be drawn 
between species occurring in New Guinea 
compared with those present in Australia. 

In several instances (C. australis, C. cloelia, C. 
comuta, C. cybele, C. dahli), the same species of 
Cloacina are found in Australia and in New 
Guinea, a phenomenon explicable in terms of host 
distribution. C. australis and C. comuta are both 
parasites of M. agilis, a wallaby which is common 
in northern Australia (Strahan 1995) and which 
invaded southern New Guinea relatively recently 
from Australia (Flannery 1995). Similarly, C. 
cloelia, C. cybele and C. dahli occur in pademelons 
of the genus Thylogale, and at least one species, T. 
stigmatica, is thought to be a recent arrival in New 
Guinea (Winter 1 997). However, the occurrence of 
these same nematodes in species of Thylogale 
restricted to New Guinea suggests that they have 
been present for a longer period of time than the 
recent invasion of T. stigmatica. 

The remaining species of Cloacina currently 
described from New Guinea occur in hosts which 
are endemic. Species of scrub wallabies Dorcopsis 
and Dorcopsulus do not occur in Australia, while 
the tree kangaroos, Dendrolagus spp., have 
apparently radiated in New Guinea and invaded 
north-eastern Australia secondarily (Flannery et al. 
1996). The recent arrival of the genus 
Dendrolagus in Australia may have contributed to 
the absence of species of Cloacina in D. lumholtzi 
and D. bennettianus. 



While the records of species of Cloacina from 
New Guinea are still fragmentary, it appears that a 
substantial New Guinean fauna exists, and its 
relationships with its hosts and the comparisons 
that can be made with Australian representatives 
may provide insights into the evolution and hence 
the diversity of this nematode genus. 

Key to species of Cloacina in macropodids 
from Papua new Guinea 

1. — Denticles present in preneural region of 

oesophageal lumen 2 

— Denticles absent 13 

2. — Paired ventral oesophageal denticles 

anterior to dorsal denticle C. dahli 

— Single dorsal denticle, or three denticles at 
same level in oesophagus 3 

3. — Anterior part of oesophagus lined with 

sclerotised bosses 4 

— Anterior oesophagus without sclerotised 
bosses 8 

4. — Anterior oesophageal bosses much larger 

than other bosses; deirid anterior to nerve 
ring; female tail swollen C. australis 

— Oesophageal bosses of similar size; deirid 
at level of nerve ring or posterior to it; 
female tail not swollen 5 

5. — Buccal capsule symmetrical in lateral view 

C. erigone 

— Buccal capsule asymmetrical in lateral 
view, more shallow on dorsal aspect ... 6 

6. — Buccal capsule prominently arched 

anteriorly C. hyperea 

— Buccal capsule not prominently arched 
anteriorly 7 

7. — Spicules 1.30-1.58 mm; distal segment of 

submedian papilla less than 1/4 length of 
proximal segment C. procris 

— Spicules 1 .67-2.07 mm; distal segment of 
submedian papilla only slightly shorter 
than proximal segment C. sterope 

8. — Dorsal denticle and 2 vestigial subventral 

denticles present 9 

— Single dorsal denticle present 10 

9. — Spicules 3.57-3.98 mm C. eurynome 

— Spicules 2.29-2.63 mm C. theope 

10. — Prominent lips present C. oweni 

— Lips absent 11 

11. — Deiridatlevelofnervering;anteriormargin 



CLOACINA FROM PAPUA NEW GUINEA 



31 



of buccal capsule arched C. sciron 

— Deirid in anterior oesophageal region, 
anterior margin of buccal capsule 
horizontal 12 

12. — Cervical cuticle and female tail inflated, 

distal segment of submedian papilla almost 
as long as proximal segment, spicules 
1.38-1.62 mm C. comma 

— Cervical cuticle and female tail not inflated, 
distal segment of submedian cephalic 
papilla much shorter than proximal 

segment, spicules 1.84-2.26 mm 

C. papuensis 

13. — Anteriormarginofbuccalcapsuleundulate 

... 14 

— Anterior margin of buccal capsule not 
undulate 18 

14. — Amphidsonorformingconicalelevations 

projecling beyond cephalic collar 1 5 

— Amphids embedded within cephalic collar 
, 16 

15. — Head deviated dorsally, cephalic collar 

elevated between adjacent submedian 
papillae C polymela 

— Head directed anteriorly, cephalic collar 
depressed between adjacent submedian 
papillae C. sapplw 

16. — External branchlets of dorsal ray arise 

before major bifurcation C, syphax 

— External branchlets of dorsal ray arise 
after major bifurcation 17 

17. — Cervical cuticle prominently inflated, 

submedian papillae project prominently 
beyond cephalic collar C. salon 

— Cervical euliele not inflated, submedian 
papillae very small, barely project above 
cephalic collar , C. solymiis 

IS. — External branchlets of dorsal ray arise 
before major bifurcation 19 

— External branchlets of dorsal ray arise 
after major bifurcation 21 



22. — 



23. 



19. — Deirid anterior to nerve ring 

C. cwictabunda 

— Deirid posterior to nerve ring 20 

20 — Spicule length 2.34-2.97 mm 

C. crethfis 

— Spicule length 1.23-1. 45 mm . 

• C. caballeroi 

21. — Buccal capsule sinuous in apical views, 
appearing as extra thickenings of wall in 

lateral or dorso-venlral views 

, C, sancus 

— Buccal capsule nol sinuous in apical views 
22 

Buccal capsule prominently arched 

anteriorly C. nephele 

Buccal capsule not prominently arched . 
23 

Amphids posterior to anterior margin of 
buccal capsule C. praxithea 

— Amphids anterior to anterior margin of 
buccal capsule 24 

24. — Oesophagus with sclerotised bosses.. 

spicule tip foot-shaped C. ctoctia 

— Oesophagus lacking bosses, spicule tip 
simple 25 

25. — Spicule length 3.90-5.05 mm, spicule tip 

simple C. hecalc 

— Spicule length 2.30-2.90 mm, spicule lip 
with enlarged flange, ala terminating 

abruptly anterior to spicule tip 

C. cybele 

Acknowledgments 

Sincere thanks are due to Tim Flannery, Terry 
Reardon, Ian Redmond, Jocrg Mayer and Ifor Owen for 
collecting the material, sometimes under Irving 
circumstances an which this paper is based and to 
Eileen Harris and Janet Martin for making the material 
available lor study Rosemary Harrigan is thanked for 
excellent technical assistance and Dave Spratl Tor 
comments on a draft of the manuscript. 



RlTlRrjNCtS 



BEVER1DGE. I. 1987. The systematic status of 
Australian Strongyloidea (Ncmatoda). Bulletin ilu 
Museum National d'Hisloire Naturelle, Paris, 41mc 
sene 9: 107-126. 

BEVER1DGE, I. 1 998. Taxonomic revision of the genus 
Cloacina von Linstow (Ncmatoda: Strongyloidea) 



from maeropodid marsupials. Invertebrate Zoology 
12: 273-508. 

BF.VERIDGE, 1. 1999. New species of Cloacina von 
Linstow, J 898 (Nematoda: Strongyloidea) parasitic 
in the stomach of the quokka, Setonix. hrachyurus 
(Marsupialia: Macropodidae), from Western 



32 



I. BEVERIDGE 



Australia. Transactions of the Royal Society of South 
Australia 123: 17-30. 

BEVERIDGE, t. & SPEARE, R. 1999. New species of 
parasitic nematodes from Dorcopsulus vanheurni 
(Marsupialia: Macropodidae) from Papua New 
Guinea. Transactions of the Royal Society of South 
Australia 123: 85-100. 

BEVERIDGE, I.. CHILTON, N. B., JOHNSON, P. M., 
SPEARE, R., SMALES, L. R. & SPRATT, D. M. 
1998. Helminth parasite communities of kangaroos 
and wallabies (Macropus spp. and Wallahia hicolor) 
from north and central Queensland. Australian 
Journal of Zoology 46: 473-^95. 

FLANNERY, T. F. 1995. 'Mammals of New Guinea'. 
Reed Books: New South Wales. 

FLANNERY, T. F., BOEAD1, & SZALAY, A. L. 1995. 
A new tree kangaroo (Dendrolagus: Marsupialia) 
from Irian Jaya, Indonesia, with notes on 
ethnography and the evolution of tree kangaroos. 
Mammalia 59: 65-84. 

FLANNERY, T. F, MARTIN, R. & SZALAY, A. L. 
1996. 'Tree kangaroos: a curious natural history'. 
Reed Books: Melbourne. 



LEW1N, R. A. 1999. 'Merdc. Excursions into scientific, 
cultural and sociohislorical coprology'. Aurum Press: 
London. 

MAWSON, P. M. 1977. Cloacina comma (Davey & 
Wood) and C. caballeroi sp. nov. (Nematoda : 
Cloacininae) from macropods from Papua. Excerta 
Parasitologica en memoria del Doctor Eduardo 
Cahallero y Cahallero. Institute de Biologia, 
Puhlicaciones Especiales 4: 455-458. 

SPEARE, R., BEVERIDGE, I. & JOHNSON, P. M. 
1983. Parasites of the agile wallaby, Macropus agilis 
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96. 

SPRATT, D. M., BEVERIDGE. I. & WALTER, E. L. 
1991. A catalogue of Australasian monotremes and 
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Records of the South Australian Museum, 
Monograph Series, 1: 1-105. 

STRAHAN, R. 1995. 'The mammals of Australia'. 
Reed Books, Chatswood. 

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DARWIN FORMATION (EARLY CRETACEOUS, NORTHERN 
TERRITORY) MARINE REPTILE REMAINS IN THE SOUTH 

AUSTRALIAN MUSEUM 

Benjamin P. Rear 



Summary 

Marine reptile remains from Early Cretaceous (Aptian/Albian) deposits of the Darwin Formation, 
Northern Territory, are described from the collections of the South Australian Museum. The 
material includes vertebra and limb girdle elements from ichthyosaurs and plesiosauroids, as well as 
the first described remains from the Northern Territory. The pliosaurid specimen (a single femur) is 
attributed to the small-bodied (up to 3 m in length) freshwater/near-shore marine taxon 
Leptocleidus, supporting interpretation of the Darwin Formation as representing a near-shore 
marine depositional environment. A comparison of the Darwin Formation marine reptile fauna with 
those known from Early Cretaceous deposits elsewhere in Australia, indicates greatest 
compositional similarity to the Hauterivian-Barremian Birdrong Sandstone fauna of Western 
Australia. A potential link between the distribution of some marine reptile taxa (eg pliosaurids) and 
the prevailing depositional environment is also suggested. 



DARWIN FORMATION (EARLY CRETACEOUS, NORTHERN TERRITORY) MARINE 
REPTILE REMAINS IN THE SOUTH AUSTRALIAN MUSEUM 



BENJAMIN P. KEAR 



REAR, B P 2002. Darwin Formation (Early Cretaceous. Northern Territory) marine icpulc 
remains in the South Australian Museum. Records »/% South Australian Museum 35(1): JJ47 

Marine reptile remains from Early Oetaceous (Aplian/Albian) deposits of the Darwin 
Formation, Northern Territory, are described from the collections of the South Australian 
Museum The material includes vertebra and limb girdle elements from ichthyosaurs and 
plesiosauroids, as well as the first described pliosaurid remains from die Northern Territory. The 
pliosaurid specimen (a single femur) is attributed to the small bodied (up to 3 m in length) 
frcshwatcr/ne.ar-shore marine taxon l.epivcleidus, supporting interpretation of the Darwin 
Formation as representing a near-shore marine dcpositional environment. A comparison of the 
Darwin Formation marine reptile fauna with those known from Early Cretaceous deposits 
elsewhere Id Austialia, indicates greatest compositional similarity to the Hauterivian-Barremian 
Hiidrong Sandstone fauna of Western Australia. A potential link between the distribution of some 
marine reptile urxa (eg pliosaurids) and die prevailing depositional environment is also suggested. 

Benjamin P. Kear (kearbenfe'saugovsagovau], *Soulh Australian Museum. North Terrace, 
Adelaide, South Australia S000; and Vertebrate Palaeontology Laboratory. School of 
Biological Sciences. University of New South Wales, UNSW Sydney. New South Wales. 
2052, 'address lor correspondence. Revised manuscript received 1 February 2002. 



Sometime in the 1960s (exact date unknown), a 
small collection of fossil marine reptile remains 
from the Early Cretaceous (Aplian/Alhian) 
deposits of Casuarina Beach, Darwin Formation, 
Northern Territory (see Murray 1987, Fig I . p. 95 
for locality map) was donated to the South 
Australian Museum. In 1994 several additional 
Darwin formation marine reptile specimens were 
acquired as part of an exchange with the Museum 
and Art Gallery of the Northern Territory A 
recent examination of these remains, which 
include isolated elements from both ichthyosaurs 
and plesiosauroids, has also revealed the presence 
of i pliosaurid femur This specimen is 
attributable to the genus Leptocteidus Andrews, 
1922 and represents the first documented pliosuur 
material front the Northern Territory. This paper 
describes Darwin Formation marine reptile 
remains currently housed in the South Australian 
Museum and provides a brief palaeoeeological 
analysts based on faunal comparison with other 
known Early Cretaceous marine replilt? bearing 
deposits. 

Marine reptile remains were first reported from 
the Darwin Formation by an anonymous author 
(1924) who described a fragmentary ichthyosaur 
skeleton from Fannie Bay near Darwin. This 
specimen was later attributed to Ichthyosaurus 
aitsrrulis (Tiechert & Mathcson 1944), however. 



Murray (1985) re-assigned the material, alonj; 
with other ichthyosaur remains from Nightcliff, 
north of Fannie Bay, to Icbthyosauria gen. et sp. 
indct. A more recent examination by Wade ( 1990| 
suggested that all ichthyosaur material from the 
Darwin Formation might be synonymous with 
flatyp ten giits longmani from the Aptian-Albian 
of Queensland. Murray (1987) described isoialed 
ichthyosaur and elasmosaurid remains from 
Casuarina Beach, assigning the material In 
Platy[>terygius sp. and Elasmosauridae gen. et sp 
indct., respectively. The presence of elasmosaurid 
and pliosaurid remains in the Casuarina Beach 
deposits was also noted by Cniickshank et al. 
(1999) but no descriptions were provided. 

A comparison of the Darwin Formation manne 
reptile fauna with those known from Early 
Cretaceous deltaic- marine deposils elsewhere in 
Auslralia (Wallumbilla Formation [Doncaster 
Member], Allaru Mudstone, Toolebuc Formation 
and Mackunda Formation, Queensland; Bulldog 
Shale, South Australia; Wallumbilla Formation 
[Doncaster Member], New South Wales, Birdrong 
Sandstone and Barrow Group, Western Australia) 
indicates greatest compositional similarity to the 
Hauterivian-Barremian Birdrong Sandstone 
fauna, with ichthyosaurs, elasmosaurids and the 
small-bodied pliosaurid Leptacleidus represented. 
A potential link between the distribution of some 



34 



IV P. KEAR 



marine rcpiile taxa (eg pliosaurids) and the 
prevailing dcpositional environment is also 
suggested The presence of LepUaeheuitts, a 
pliosaund genus known otherwise from 
freshwater and near-shore marine deposits 
(Andrews 191 1 , 1922; Stromer 1935; Cruiekshank 
1997; Cruiekshank & Long 1997; Cruiekshank et 
al. 1999) supports interpretation of the Darwin 
Formation as representing a shallow near-shore 
marine dcpositional environment (sensu Smart & 
Senior 1980; Murray 1985; Henderson 1998). 



Materials And Methods 

Institutional abbreviations used. SAM, South 
Australian Museum, Adelaide; N'I'M, Museum 
and Art Gallery of the Northern Territory, Darwin; 
AM, Australian Museum. Sydney 

Lilhostratigraphic nomenclature follows Mory 
(1988) for Early Cretaceous deposits of the 
Darwin area and Burton & Mason (1998) for the 
opal-bearing sediments of White Cliffs, New 
South Wales Systematic terminology follows 
Montani (1999) for Ichlhyoptcrygia and Brown 
(1981) for Plcsiosauria except in the subdivision 
of Pliosauroidea. in which Polycolylidae is 
retained as a separate family (sensu Williston 
1925; While 1940; Pcrsson 1963; Welles 1962; 
Carpenter 1996, 1997). Functional subdivision of 
the vertehral column in ichihyopterygians follows 
Buchholtz (2001). with designation of structural 
units as neck, trunk, tail stock and fluke. 
Interpretation of plesiosaur pelvic and femoral 
musculature follows Robinson (1975) and 
Lingham-Soliar (2000). All measurements were 
taken using callipers and arc in millimetres (mm). 



Geological Sftdng And Prlsiirvation 

The marine reptile-bearing deposits of the 
Darwin Formation are typically characterised by 
glaueonitk sandstone and radiolarian mudstnne 
with basal quartzose conglomerate and localised 
nodular phosphorite horizons (Henderson 1998). 
The unit forms part of the BaihUT.st Island Group, 
which rests nonconformably on Precambrian 
basement within the Money Shoals Platform (see 
Fig. 1. p. 117 and Fig. 6. p. 124 of Henderson 
|§98 for locality and straugraphic selling). At its 
base the Balhurst Island Group is composed of 
the laterally equivalent Darwin Formation and 
Marligar Formation, which are overlain in turn by 
ihc Wangarlu Mudstone and Moonkinu 



Formation. 

The Darwin Formation has historically been 
considered Albion in age on the basis of its 
macroinvertebrate (Day 1969; Skwarko 1966, 
1968; Henderson 1990) and palynomorph 
(Morgan 1980) assemblages, however, more 
recent analyses (Henderson 1998; D Meginan 
pers. comm. 2000) suggest an upper Aptian age. 
The Marligat Formation has been dated as 
Ntocomian/Aptian by palynological studies 
(Burger 1978) and upper Aptian because of its 
laterally equivalent relationship with the Darwin 
Formation (Henderson 1998). The overlying 
Wangarlu Mudstone is regarded us upper Albian- 
Cenomanian (possibly wilh a basal lower Alhian 
component, Henderson 1998) on the basis of 
ammonite (Henderson 1990) and palynologieal 
(Norvick & Burger 1975, Burger 1978) data. The 
uppermost Moonkinu Formation is considered 
Cenomanian-Turonian on account of its 
ammonite (Wright 1963) and bivalve/gastropod/ 
scaphopod (Skwarko 1983) faunas. 

The glauconitic sandstones and predominant 
radiolarian mudslones of Ihe Darwin Formation 
reflect a near-shore marine/para lie dcpositional 
environment (Smart & Senior 1980; Henderson 
1998). The presence of a limited benthic 
invertebrate fauna also suggests unfavourable, 
possibly anoxic bottom conditions (Henderson 
1998). 

The Darwin Formation marine reptile remains 
generally occur as heavily weathered isolated 
elements, although several fragmentary skeletons 
have been recovered (Murray 1985, D Megirian 
pers. comm. 2000). The disarticulated nature of 
the remains led Murray (1985) to speculate that an 
inshore and/or tidal depositional environment may 
have facilitated stranding of decaying carcasses 
on mudflats prior to burial. 



Systematic* 

D1APSIDA Osbom, 1903 

ICHTHYOPTERYGIA Owen, 1860 

ICHTHYOSAURtA de Btainville, 1S35 

OPHTHAI.MOSAURIA Appleby, 1956 (scn.w 
Montani 1999) 



PkUypHrygius von Huene, I'm 
cf. Platyprerygius luiigmani 



DARWIN FORMATION MARINE REPTILE REMAINS 



35 




FIGURE 1- SAM P35039, cf. Plaiprerygius hiMgmtml dorsal vertebra in (A) anterior and (B) lateral views. Scale 
bar is 40 mm. 



Material 

SAM P35039 (Figs 1A, B; Tabic I), four 
associated mid-posterior trunk vertebral centra; 
SAM P35426 (Table I), two articulated anterior 



trunk centra; SAM P35427 (Table 1), three 
associated anterior tail stock centra; SAM P35429 
(Table 1), two associated sections of eight and 
four articulated anterior? tail stock centra; SAM 




FIGURE 2. SAM P35430. cf Plntpterygius h/ngmuni caudal vertebra in (A) anterior, (R) lateral, (C) ventral and 
(D) dorsal views. Scale bar is 20 mm. 



36 



B. P KEAR 



TABLE 1. Measurements (mm) of cf Plotyprerygius 
longmani vertebral centra from the Darwin Formation, 
Northern Territory. 



Specimen 


Length 


Width 


Height 


SAM P35039 


39.6 


91.2 


98.9 




33.1 


88.9 


90.2 




30.2 


80.1 


89.2 




32,3 


62.6 


66.1 


SAMP35426 


40.6 


85.5 


88.9 




37.0 


86 1 


89.7 


SAM P35427 


40.7 


94.0 


96.4 




30.1 


86.2 


92.4 




25 K 


102.8 


109.6 


♦SAM P35429 


24 V 


81.6 


69.9 




25.8 


83.1 


7S.2 




26.4 


86.4 


79 1 




25.2 


648 


83.9 




20.5 


- 


78,2 




21.6 


58.3 


60.1 




2t.l 


58.9 


50.8 




22.3 


- 


57,2 




174 


- 


55 1 




18.9 


- 


46.6 




14.1 


- 


469 


SAM P35430 


20.8 


68.9 


59.7 




20,8 


61 1 


62 J 




18.7 


55 7 


56.J 




14.6 


53.2 


50.4 




15.3 


<»7,9 


48.3 




15,3 


42.8 


42.6 




152 


43.6 


45.0 




129 


41.5 


41.3 



some centra represented by fragments only 



P35430 (Figs 2A, B, C, D; Table 1), nine 
associated anterior tail stock centra (juvenile). 

Locality 

Casuarina Beach, north of Darwin. Northern 
Territory. Darwin Formation, Aptian or Albian. 

Description 

All of the preserved centra are disc-like and 
anteropostcriorly compressed, with the dorsal 
length subequal to the ventral. Tail stock centra 
(Figs 2A, B, C, D) exhibit a more marked degree 
of anteroposterior compression relative to those 
from the trunk region (Figs I A. B) and arc slightly 



ovoid in outline, with height being approximately 
1.5 limes the transverse width. The articular 
surfaces of alJ centra are deeply amphicoelous, 
with a distinct central notochordal pit. The neural 
canal is generally broad and shallow, becoming 
narrower in centra from the tail stock region. The 
neural arch facets are narrow and rectangular in 
all centra. Where preserved, both the diapophyses 
and parapophyses of the trunk centra are distinctly 
raised and subcircular. This is unlike centra from 
the tail stock region, in which only the ovoid 
diapophysis is present and positioned low on the 
lateral centrum surface. The ventral surfaces of at 
least two of the recovered centra from the tail 
stock region show distinct haemal arch facets. 
These are slightly concave and saddle-shaped in 
outline with low ridge-like margins. Medially, the 
haemal arch facets border the very weakly 
concave, spool-shaped depression for the haemal 
canal. 

Remarks 

All of the ichthyosaur vertebral centra examined 
can be distinguished from those of basal 
ichthyopterygians by their disc-like shape and 
strong anteroposterior compression (the more 
plcsiomorphic Grippidia having cylindrical centra, 
Montani 1999). The absence of any further 
diagnostic features restricts attribution of the 
specimens to Ichthyosauria gen. et sp. indet. 
However, tentative assignment of the remains In 
cf. Plaryplerygius longmani is justifiable on the 
basis of- 1) overall similarity to /'. longmani 
vertebral material described from ihc A Dam 
Mudstone and Toolebuc Formation, Queensland 
(Wade 1990) and Darwin Formation (Murray 
1985. 1987 sensu Wade 1990); and 2) current 
recognition of P. longmani as the only valid 
ichthyosaur species from the Aptian-Albian of 
Australia (Wade 1990; Bardet et al. 1994, 
Arkhangelsky 1998; Montani 1999). 

Buchholtz (2001) discussed differentiation of 
structural units within the vertebral column oT 
Jurassic ichthyosaurs with relation to variation in 
swimming styles between taxa. Following on from 
this study, it can be suggested that differences 
observed in the Darwin Formation centra (relating 
to rcgionaiiscd morphological variation along Ihe 
vertebral column) can be used to predict preferred 
swimming style in the Cretaceous P. longmani. 
The marked increase in height proportionate to 
width, and anteroposterior compression of the tail 
slock relative lo trunk centra, in the Darwin 
Formation specimens is comparable to the 
condition recorded in derived thunnilonn-model 



DARWIN FORMATION MARINE REPTILE REMAINS 



37 



ichthyosaurs such as Ophthalmosaurus icenicus 
and Stenopterygius quadrissicus (Buchhotli 
2001). Wade (1990) and Broili (1907) noted that 
the anterior fluke centra in P. longmani and P. 
plarsductylus were distinctly higher lhan wide, a 
feature also conforming to the predicted 
thunnilorm condition. Taken together therefore, 
these characteristics may suggest that 
Plotypterygius spp. employed an axial oscillatory 
swimming mode and probahly shared many other 
key Irails (see Montani 1999: Buchholtz 2001) 
with derived thunni form-model ichthyosaurs. 



Material 

SAM P35431 (Figs 3A, B, C), incomplete and 
badly weathered element tentatively identified as 
Ihe ventral plate portion of a left ischium. 
Orientation is based on position of the lateral 
margin and its altitude relative to the horizontal 
axis: SAM P35434 (Figs 4A, B, C), right ischium 
lacking part of antcromedial margin. 

Locality 

Casuarina Beach, north of Darwin, Northern 
Territory. Darwin Formation. Aptian or Albian. 



SAUROPTERYG1A Owen, I860 
PLESIOSAURIA de Blainville, 1835 
PLESIOSAUROIDEA (Gray, 1825) Welles. 1943 
Plesiosauroidea gen. et sp indel 



Description 

Dimensions of remains are: SAM P35431, total 
length of 96.7 mm; total width is 98.9 mm; 
maximum height is lateral extremity is 22.5 mm; 
maximum length of lateral extremity is 55.3 mm; 
maximum height of ventral plate is 45.9 mm 
SAM P35434, tolal length is 74.6 mm; total width 




FIGURE 3. SAM P3543I, Plesiosauroidea gen. et sp. indct. partial ischium in (A) dorsal, (B) medial and (C) 
posterior views. Scale bar is 40 mm. 



38 



B. P KEAR 




FIGURE 4. SAM P35434, Plesiosauroidea gen. et sp. indct. ischium in (A) dorsal, (B) posterior and (C) lateral 
views. Scale bar is 40 mm 



is 97.4 mm; maximum height of the of lateral 
articular surface is 52. 1 mm; maximum length of 
lateral articular surface is 46.1 mm; maximum 
height of ventral plate is 10.8 mm. Both SAM 
P35431 and SAM P35434 are halchcl-shapcd in 
dorsal outline with an anteroposteriorly expanded 
ventral plate. The lateral articular extremity of 
SAM P35434 is massive but has been lost in SAM 
P35431. Medially, the ventral plate portion of 
both elements is dorsovenlrally compressed with 
an inclined medial edge for contact with the 
opposing ischium. This surface is oriented 
approximately 10° ventral to a horizontal plane 
through the lateral articular extremity, and is 
deeply pitted, suggesting the presence of cartilage. 
Both the dorsal and ventral surfaces of the ventral 



plate in SAM P35434 are rugose, possibly for 
attachment of the m. puboischiofemoralis inlernus 
and m. p. cxtemus. respectively. A weak ridge on 
the posteromedial margin of the ventral plate may 
also have supported slips of the m. 
ischiolrochantcricus. The ventral plate of both 
specimens is not greatly anteroposteriorly 
expanded (with total length being just slighlly less 
than the total transverse width of each element) 
and is separated from the lateral articular 
extremity by a constricted waist-like midsection. 
This is ovoid to circular in cross-section, 
becoming increasingly dorsovenlrally thickened 
towards the lateral articular surface (nol preserved 
in SAM P3543I). The articular surface itselT is 
convex, with the rectangular anterior pubis facet 



DARWIN FORMATION MARINE REPTILE REMAINS 



39 



uffsel approximately 195" relative to the 
longitudinal axis of the articular surface. The 
pubis facet is separated posteriorly from the 
rectangular median acetabular facet, and in turn 
from lh<* lobatc posterior-most ilial facet (offset 
approximately 150" relative to the longitudinal 
axis of the articular surface) by weak parallel 
ridges. The articular sutface margin is produced 
into a crennate rim, which is continuous ventrally, 
with a low rugose tuberosity possibly marking part 
of the ra. puboischiofenioralis internus 
attachment. 

Remarks 

Despite being heavily weathered, SAM 
P35431 is identified as the ventral plate portion 
of a plesiosaurian left ischium on the basis of its 
hatchet-shaped outline and general morphology. 
Nevertheless, the specimen is unusual in its 
markedly dorso ventrally deep, medial articular 
margin. SAM P3543I was initially registered as 
an ichthyosaurian humerus belonging to 
Platvpterygius sp. However, such an 
identification is unlikely as the humeri of 
tlatyfiicrygius spp. arc much more robust, with 
rectangular to spool-shaped dorsal outline and 
prominent ridge-like tuberosities on both the 
dorsal and ventral surfaces (eg Kiprijanoff 1881; 
Bioih 1907; Kuhn 1946; McGowan 1972: Wade 
1984; Murray 1987; Choo 1999). SAM P35431 
also differs from the larger girdle elements of 
Pluiyi'terygius spp., in which the coracoid is 
subcircular in outline with dorso ventrally deep 
glenuid/intercoracoid facets (Broili 1907; Wade 
1984); and both the scapula and ischiopubis are 
strap- like with predominantly narrow, ridge-like 
angular surfaces (Hroili 1907; Nace 1939; 
Wuiiay 1987). 

SAM P35434 probahly represents a juvenile 
individual because of its small size and poor 
ossification of the articular surfaces. Both it and 
SAM P35431 can be assigned to Plesiosauroidea 
on the basis of their anteroposteriorly short ventral 
plates^ This differs from the condition in 
pliosauroids in which the ventral plate of the 
ischium bears a marked posterior expansion (Mehl 
1912; Andrews 1913; Tarlo 1959. 1960; Brown 
198 1). The presence of a relatively weak posterior 
expansion in Eurycleidus (Cmickshank 1994) and 
Lepiotteidus (pers. nbs, AM F99374 I ram Ihe 
Apiian-Albian Bulldog Shale of CooheT Pedy, 
South Australia), however, suggests that this 
character state may not be universal for the group. 
SAM P35434 is unusual in its proportionately 
massive articular head relative to the ventral plate. 



The phylogenetic significance of this feature, if 
any, is uncertain because of its immature stage of 
development. 



ELASMOSAURIDAE Cope, 1869 

Elasmosauridae gen. ct sp. indel. 

Material 
SAM P35432 (Figs 5 A, B, C, D), a single 

weathered posterior cervical vertebral centrum. 

Locality 

Casuarina Beach, north of Darwin, Northern 
Territory. Darwin Formation, Aptian or Albian. 

Description 

The surface of the centrum is badly weathered 
with very little periosteal bone remaining. Total 
lenglh is 72.8 mm, width of anterior surface is 
109.5 mm; depth of anterior surface is 85.2 mm 
In general proportions, the centrum is 
anteroposteriorly short relative to both the height 
and width of the anterior articuUr surface. Both 
the lateral and ventral surfaces are shallowly 
concave, with the ventral surface being pierced by 
two large foramina. The articular surfaces arc 
poorly preserved but clearly platycoelous, with a 
raised convex rim. The neural arch facets are 
indistinct but prominent bosses high on the lateral 
surface of the centrum indicate positioning of the 
rib facets. 

Remarks 

SAM P35432 is tentatively assigned to 
Elasmosauridae on the basis of its platycoelous 
articular surfaces. Murray (1987) reported an 
elasmosaurid cervical with very shallowly concave 
articular facets from the Casuarina Beach locality. 
Brown (1981, 1993) and Bardet et al. (1999) 
regarded the presence of platycoelous articular 
surfaces on the cervical centra to be a potential 
synapomorphy for Elasmosauridae. The character 
state has also been recorded in the Maastrichtian 
cryploclidid Monurneria (Chattergee & Small 
1989), though Bardet et al. (1991) suggested that 
this taxon may represent a derived elasmosaurid. 

The proportionately short centrum length 
relative to height of SAM P35432 differs from the 
common condition in elasmosaurids (such as that 
described by Murray 1987), in which the cervical 
centra are markedly elongate (Brown 1993; Bardet 
et al. 1999), The presence of relatively short 
cervical centra is considered plesiomorphic among 



40 



B P. KEAR 




FIGURE 5. SAM P35432. Elasmosaurjdae gen. el sp. indel. posterior cervical centrum in (A) dorsal. (B) ventral, 
<C) anterior and (D) lateral views. Scale bar is 40 mm. 



plesiosauroids iBrown 1993; Bardet et al. 1909). 
However, the feature has been described in the 
potential elasmosaurid Ut\a (sensu Bardet et al 
1991) Aristonectes (Cabrera 1941) and 
Mnrturneria (Chattergee & Small 1989) as well 
as in indeterminate elasmosaurid posterior 
cervicals from unspecified (probably Albian) 
deposits near Oodnadatta (Freytag 1964) and Ihe 
Neales River region (SAM P6181, Persson 1960). 
South Australia; Toolebuc Formation, Queensland 
(Kear 2001); and Molccap Grccnsand, Western 
Australia (Long & Cruickshank 1998). The 
presence of proportionately elongate cervicals 
may therefore not necessarily be universal 
amongst elasmosaurids, with some taxa exhibiting 
relatively short centra, particularly in the posterior 
cervical region. 

SAM P35432 is notable for its possession of a 
raised convex rim surrounding the central 



concavity of the articular surface. Brown et al. 
(1986) and Brown (1993) considered this a 
plesiomorphic condition commonly associated 
with members of ihe Cryptoclididae and 
Plesiosaundae. Tn contrast, elasmosaurids, 
including basal taxa such as Occiianosaurui 
(Bardet el al. 1999) and Mtiraennsaurua (Andrews 
1910, Brown 1981, 1993), generally exhibit a 
more open V- shape, in which the articular surface 
margins form an abrupt angle with the centrum 
sides, A similar trait is evident in elasmosaurid 
material from the Toolebuc Formation of 
Queensland (Kear 2001 ), and is indicated (hut not 
described) in the figured Darwin Foimaiion 
specimen (NTM P8727-70, Fig. I, p. 97) of 
Murray (1987). Carpenter (1996), however, 
considered articular nm morphology to be a 
variable character amongst plesiosaurs and of 
questionable diagnostic value. 



DARWIN FORMA I It >N MARINE REPTILE REMAINS 



41 



PLIOSAUROIDEA Seeley. 1874 
PLIOSAURIDAE Seeley, 1874 
Leptocteidus Andrews, 1022 
Leptocteidus sp. 



Mmenul 

SAM P35053, cast of NTM P9 1.3-5 (Figs 6A, 
B, C) nghi femui. 

Locality 

Casuanna Beach, north of Darwin, Northern 
Territory. Darwin Formation, Apt ran or Albian. 

Inscription 

A near complete right lemur lacking pari Of 
postcrodisial margin Total length is 260 8 mm, 




FIGURE 6. SAM P35053 (cast or NTM P913-5). 
Leptocleidus sp. femur in (A) anterior, (B) dorsal and 
(C) proximal views. Scale bar is 40 mm. 



tolal proximal length is 47.9 rnm; total proximal 
height (including trochanter) is 51.5 mm; width 
across base of dorsal trochanter is 37.2 mm. total 
distal length (not including missing posterior 
margin) ts 91.6 mm, total distal height is 
25.9 mm. The anterior edge of the femur is weakly 
sigmoidal. wilh a strongly convex distal extremity. 
The posterior edge is uniformly concave along its 
entire length. Both the dorsal and ventral surfaces 
are shal lowly concave along (heir longitudinal 
axis, however, the ventral surface becomes 
convex and upturned towards the distal extremity. 
Proximatly, the femur is greatly dorsovenlrally 
expanded and separated from its distal section by 
a constricted neck. The dorsal trochanter is robust 
and anteroposterior^ constricted at its base. It i 
separated from the ovoid glenoid articular surface 
by a shallow groove. The surfaces of both the 
dorsal Irochanler and glenoid articulation are 
deeply pitted, suggesting the presence of cartilage. 
The femoral shaft is weakly anteroposterior^ 
constricted and ovoid in cross-sectaiu, with it* 
posterior margin bearing a raised rugose 
tuberosity, possibly for attachment of the m. 
caudifemoralis. A second low tuberosity is present 
on the median ventral surface and may represent 
pan of the m. puboischiofemoralis insertion. The 
distal femoral extremity is fan-shaped in 
dorsovential outline, with a weakly projecting 
anterodistal margin. The distal articular surface is 
narrow and elliptical, with a raised median ndge 
separating the very large radial facet from the 
remains of the ulnar facet. 

Remark, x 

Despite the poor existing knowledge of femoral 
morphology in Leptocteidus spp., SAM P35053 
(NTM P913-5) can be readily distinguished from 
the proportionately shorter and stouter propodtaJs 
of elasmosaunds (Brown 1981, Murray 1987) by 
its anteropos(eriorly broad, flat femoral shaft, 
greatly expanded distal extremity and sigmoidal 
anterior margin, These characteristics are very 
similar to those of the fragmentary femora of L. 
capensis (Andrews 1911) and well-preserved 
material (AM F99374. SAM P15980) from the 
Bulldog Shale of South Australia (Figs 7A, B). 
The femur of L. clemai (Cruickshank & Long 
1997) differs in its possession of a concave 
anterior margin. This feature, along with a 
proportionately more slender shaft, is also present 
in the femora of polycotylids and most other 
pliosauroids (Andrews 1913; Welles 1943. 1962; 
Brown 1981; Storrs 1999). A sigmoidal anterior 
femoral and humeral margin has, however, been 



42 



B P. KEAR 



TABLE 2. Marine reptile groups from Early Cretaceous deltaic-marine deposits of Australia. Pliosauroids are 
separated into small-bodied (eg Lepiocleidus) and large-bodied (eg Kronosaurus) forms because of their differing 
ecological roles. The poorly known laxon Cimolmsaurus muccoyi' is recorded separately because of its unknown 
affinity. Indeterminate plesiosaur material from the marine Bungil Formation, Queensland and freshwater deposits 
of the Griman Creek Formation. New South Wales, and Strzelcki and Otway Groups, Victoria is excluded pending 
a more thorough examination of the remains. Taxa from the opal bearing deposits of White Cliffs, New South 
Wales are placed within the Wallumbilla Formation in accordance with the litlioslialigraphic nomenclature of 
Burton & Mason (1998). Source texts include; Etheridge ( 1904)'. Longman ( 1924)- Tierchert & Malbcson ( 1944)' 
Persson (I960 4 . 1%3\ 1982"), Ludbrook (196fi) 7 , Condon (1968)". Pledge (1980)'. Smart & Senior (1980)'", 
Gaffncy (1981)". Molnar (I982' : , I991 IV 1, Wade ( 19K4 H , 1990 15 ), Murray (J98S"\ 1987"). Hocking et al. (I9X7)'\ 
Burger (1988)'*', Bardet (1992) 20 , Thulborn & Turner (1993) 3 ', Kricg &. Rodgers (I995) :! , Cruickshank & Long 
<1997>'\ Henderson (1998)=', Long (1998) 25 , Long & CruiekshanMWS)*. Cruickshank ct al. (1999) 2 '. Choo 
(1999) 3 ", KearttOOJ) 1 *. 



Darwin Formation 
(Money Shoals Platform) 
Apttan/Albiun 



Ichthyosauria v " m< >" 
Elasmosauridae"" 
Small pliosaund 27 



Wallumbilla Formation 
(Eromanga Basin) 
Aptian-Albian 



Allaru Mudstone 
(Eromanga Basin) 
Albian 



Toolehuc Formation 
(hromnnga Basin) 
Albian 



lchthyosauria Ichlhyosauria' 3 n ■" li:5 

Elasmosauridac 1 ' UJ5 - ii Elasmosauridae 4 ' 512 - 1315 

Large pliosaund '■MAWM Chelonioidea' 115 
Polycotylidae' -''" 
'C. nwccovi' UM 



Ichthyosauria'-"' 1415 -" 
Elasmosauridae 4 3 * a "' : '' :!,! '' 
Large pliosaurid MMJ l) 
Polycotylidae :l ' 1 " ? 

Chclomoidea'" 21 '- 25 



Shallow near-shore 
marine 11 / paralic"?/ 
possibly tidal' 4 



Coastal / offshore shallow Shallow marine" 



marine 



10.1" 



Shallow maiinc"* 



cited as a potentially diagnostic character state for 
the polycolylid genus Folycotylus (Stons 1999). 



Discussion 

The Darwin Formation sediments have been 
interpreted as representing a shallow near-shore 
marinc/parahc (Smart & Senior 1980; Henderson 
1998) to possibly tidal (Murray 1985) deposition;!! 
environment. The presence of the small pliosaund 
taxon Lepiocleidus, known elsewhere from 
freshwater and near-shore marine deposits 
(Andrews 1911, 1922; Stromer 193.5; Cruickshank 
1997; Cruickshank & Long 1997; Cruickshank et 
al. 1999), is consistent with this interpretation. 

A comparison (Tabic 2) of the Darwin 
Formation marine reptile fauna with those known 
from Early Cretaceous deltaic-marine deposits 
elsewhere in Australia (Wallumbilla Formation 
(Doncaster Member], Allaru Mudstone, Toolehuc 
Formation and Mackunda Formation, Queensland; 
Bulldog Shale. South Australia; Wallumbilla 
Formation [Doncaster Member|. New South 
Wales; Birdrong Sandstone and Barrow Group, 



Western Australia) indicates greatest 
compositional similarity to the Hauterivian- 
Barrerman Birdrong Sandstone fauna, with 
ichthyosaurs, elasmosaimds and the small-bodied 
pliosaund Lepiocleidus represented. Throughout 
the Early Cretaceous marine deposits, 
ichthyosaurs show a wide distribution (occurring 
in all currently recognised deposits except the 
Albian Mackunda Formation and unspecified 
Berriasian subsurface sediments of the Barrow 
Group), as do elasmosaurids, which have been 
recorded from all localities except those of the 
Albian Mackunda Formation and Berriasian 
Barrow Group. 

Cruickshank et al. (1999) indicated the possible 
presence of cryptoclidids in the Aptian-Albian 
Bulldog Shale. This is supported by more recent 
observations of the South Australian fauna (Keui 
unpubl.) but as yet Ihe group has not been 
recorded from any other Australian deposit, 

The distribution of small- and large-bodied 
pLiosaurid taxa in the Australian Early Cretaceous 
is variable, with smaller forms (represented by at 
least two species of Leptorleidus, Cruickshank ct 
al. 1999) confined to the Darwin Formation. 



DARWIN FORMATION MARINE REPTILE REMAINS 



4? 



TABLE 2. (com.) 



Mackunda Formation 
(Eromanga Basin) 
Albian 



Bulldog Shale 
(Eromanga Basin) 
Aptian-Albian 



Birdrong Sandstone 
(Carnarvon Basin) 
Hauterivian-Barremian 



Barrow Group 
(Carnarvon Basin) 
Berriasian 



Polycotylidae" 



Ichthyosauria' ,l - u " 
Elasmosauridac* 27 
Small pliosaurid"- 5 " 
Large pliosaurid'' 
Cryptoclididae?" 



Ichthyosauria 115 " 18 
Elasmosauridae 2 "" 
Small pliosaurid 2 ' " 



Small pliosaurid 2 '' 



Paralic" 



Shallow marine"/ 
offshore' 



Coaslal near-shore 
shallow marine 1 "/ 
paralic 8 



Fluviatile-dcltaic 1 * 



Bulldog Shale and Birdrong Sandstone. Larger 
pliosaurid remains occur only in the Aptian- 
Albian Wallumbilla Formation (Doncaster 
Member), Albian Toolebuc Formation and 
Bulldog Shale deposits. This division may he 
linked to the different habitat preferences and 
ecological roles of small- and large-bodied 
pliosaurid taxa, wilh offshore environments 
preferentially supporting larger-bodied forms. An 
exception is seen in the shallow offshore marine 
(Ludbrook 1966; Krieg & Rogers 1995) Bulldog 
Shale, which includes both small- and large- 
bodied pliosaurid remains (the latter being 
represented by an isolated tooth, SAM P22525, 
from the opal-bearing sediments of Coobcr Pedy. 
figured by Pledge 1980, p. 8). 

Distribution of polycotylids in the shallow 
marine (Smart & Senior 1980; Burger 1988) 
Wallumbilla Formation, Toolebuc Formation and 
paralic (Smart & Senior 1980) Mackunda 
Formation may, as with large-bodied pliosaurids, 
be related to a preference for predominantly 
offshore marine environments. However, 
individuals also appear to have readily 



frequented more inshore habitats, as suggested 
by their occurrence in the Mackunda Formation 
deposits. 

Restriction of chclonioids to the upper Albian 
Allaru Mudstonc and Toolebuc Formation may be 
the product of both the group's temporal 
distribution (the earliest-known chelonioid being 
recorded from the upper Aptian - lower Albian 
Santana Formation of Brazil, Hirayama 1997, 
1998) as well as environmental factors such as 
prevailing water temperatures. Studies of 
sedimentary sequences (Frakes & Francis 1988; 
Frakcs & Krassay 1992; Frakes et al. 1995: 
Constantine et al. 1998), climatic modelling 
(Barron & Washington 1982) and isotope data 
(Gregory el al. 1989; Pirrie ei al. 1995) have 
suggested that strongly seasonal climates with 
winter freezing and at least seasonal sea ice 
characterised high latitudes in the Early 
Cretaceous of Australia. Such conditions may well 
have been unfavourable to chelotiioids and 
provided an effective barrier to the group's 
dispersal into the Australian region prior to the 
upper Albian. 



44 



B.P KEAR 




FIGURE 7. SAM PI5980 juvenile Leptocleidus sp. humerus (A) and femur (B) from Andamooka, South Australia. 
Scale bar is 20 mm. 



Acknowledgments 

I wish to thank Dirk Megirian of the Museum and 
Art Gallery of the Northern Territory for assistance 
with locating literature on the Darwin Formation 
deposits, and generous provision of NTM P913-5 for 
study. Tim Flannery, Neville Pledge. Jo Bain and Ben 
Head of the South Australian Museum provided 



support, enthusiasm, preparation facilities and 
materials. This manuscript benefited greatly from the 
comments of Dirk Megirian and Arthur Cruickshank 
of the Leicester City Museums Service, New Walk 
Museum. Origin Energy, The Advertiser, the 
Waterhouse Club and the Coober Pedy Tourism 
Association provided financial support for this 
research. 



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TENTACULARIID CESTODES OF THE ORDER TRYPANORHNCHA 
(PLATYHELMINTHES) FROM THE AUSTRALIAN REGION 

H. W. Palm & I . Beveridge 



Summary 

The present study summarises information on tentaculariid trypanorhynchs from Australian waters. 
A total of 19 species from the genera Nybelina Poche, 1926, Heteronybelinia Palm, 1999, 
Mixonybelinia Palm, 1999, Kotorella Euzet & Radujkovic, 1989 and Kotorelliella gen. nov. were 
identified: N. aequidentata (Shipley & Hornell, 1906); Nybelina africana Dollfus, 1960; N. 
hemipristis sp. nov.; N. jayapaulazariahi Reimer, 1980; N. mehlhorni sp. nov.; N. schmidti Palm, 
1999; N. scoliodoni (Vijayalakshmi, Vijayalakshmi & Gangadharam, 1996); N. strongyla Dollfus, 
1960; N. thyrsites Korotaeva, 1971; N. victoriae sp. nov.; Heteronybelinia australis sp. nov.; H. 
estigmena (Dollfus, 1960); H. pseudorobusta sp. nov.; Mixonybelinia beveridgei (palm. Walter, 
Schwerdtfeger & Reimer, 1997); M. cribbi sp. nov.; M. edwinlintoni (Dollfus, 1960); M. southwelli 
Palm & Walter, 1999; Kotorella pronosoma (Stossich, 1901) and KotorellieUa jonesi gen. et sp. 
nov. The new genus Kotorelliella is characterised by a homeocanthous, heteromorphous metabasal 
armature and a basal armature with additional interpolated hooks on the external surface of the 
tentacle, thus appealing heterocanthous. The new species appears to be closely related to Kotorella 
pronosoma. The tentaculariid trypanorhynch fauna in Australian waters is pecies rich, with 22 
(48%) of the total of 46 known species occurring in waters of the region. Eleven new locality, and 
20 new host records are reported. 



TENTACULARIID CESTODES OF THE ORDER TRYPANORHVNCHA 
(PLATYHELMINTHKS) FROM THE AUSTRALIAN REGION 

H. W. PALM & I. BEVERIDGE 

PALM. H. W. & BEVERIDGE. I. 2002. Tentaculariid ccstodes of Ihe order Trypanorhynch;, 
tPlalyhelminlhes) from die Ausuahan region. Records of the South Australian Museum 35(1): 
49-78. 

The present study summarises information on tentaculariid try panorhynchs from Austialijn 
waters. A total of 19 species from uV genera Nybelinia Pocbe, 1926, Hetcrtmshclihia Palm, 
1999. MUonybelinia Palm, 1999. Kotorella Euzet & RadujkoVic. 1989 and Kolorelhellu gen 
nov. were identified: N. aequidentata (Shipley & Hornet!, 1906); Nybelinia africatia Dollfus, 
I960: N. hemipristis sp. nov.; A*, jayapaularariahi Rcimer, 1980; N. mehllwrni sp. nov ; N. 
schnidii Palm, 1999; N. scoliodoni (Vijayalakshmi, Vijayalakshmi & Gangadharam, 1996); N. 
strongvki Dollftis. 1960: N. thyrsites Korolaeva, 1971; N. vktoriae sp. nov ; Htterotiybelinia 
ausimlis sp. nov.; H esligmena tDollfus, I960); //. pseudorohusta sp. nov.; Mixonybelinia 
beveridgri (Palm. Walter. Schwerdlfeger & Reimer, 1997): M. cribbi sp. nov., A*, edwinlimoni 
tDollfus, I960); M smtrluuflti Palm & Walter, 1999; Kotoretta pronosoma (Stossich, 1901) 
and KmnreHiella jonesi gen. el sp, nov. The new genus Ktiwrclliella is characterised by a 
homeoacanthnuK, heiemmorphous metabasal armature and a basal armature with additional 
interpolated hooks on the external surface of the tentacle thus appearing hctetoacantbous. The 
new species appears to be closely related to Kotorella pronosoma. The tentaculariid 
trypanorhynch fauna in Australian waters is species rich, with 22 (48%) of the tolal of 46 
known species occurring in waters of the region. Eleven new locality, and 20 new host records 
are reported. 

H. \V. Palm, Insliiul fur Zoomorpltologie, Zellbiologie und Parasitologic. Hcinricli-Heinc 
Universilat Dusscldorf. UnivcrsitatsstraBe I, D-40225 Dtisseldorf, Getmany, current address: 
Centre for Coastal and Marine Resource Studies, Faculty of Fisheries and Marine Sciences. 
8ogor Agricultural University. 16680 Bogor, Indonesia; email; hpalmisindo.net id, 
hpalm@gmx.net. and I. Bcvcridgc, Department of Veterinary Science, University of Melbourne, 
Parkville, Victoria 3052, Australia. Manuscript received 20 November 2000 

Palm (1999), Palm et al. (1997), Palm and unidentified species of Nybelinia were not 

Overstrcet (2000) and Palm and Walter (1999, described until 1942 (Dollfus 1942). Korotaeva 

2000) partially revised the trypanorhynch cestode (1971) named the Nybelinia species of Dollfus 

family Tentaculariidae Poche, 1926 on the basis (1942) N thyrsites and subsequently (Korotaeva 

of material deposited in museums in London 1974) reported additional unidentified Nybelinia 

Paris, Vienna and the United States. Following spp. from other Fishes from southern Australia 

the erection of two new genera, f-fetemnybelinia Lester et al (1988) and Sewell and Lester (1988.) 

Palm, 1999 and Mixonybelinia Palm, 1999 by reported Tentacuiaria sp. from orange roughy. 

Palm (1999), Palm and Walter (2000) gave a Hoplostethus atlanticus Colletl, 1889 off the 

summary Of the current state of knowledge Within coasts of South Australia and Tasmania, while 

the tentaculariid trypanorhynch genera. Thus, to Sewell and Lester (1995) found Nybelinia sp. in 

date, a total of 39 species of Kotorella, Nybelinia, jewfish, Rexea solaiulri (Cuvier & Valenciennes. 

tleteronybelinia and Miyonybelinia are considered 1832) from southern Australian waters. Beveridge 

valid (Palm &. Waller 2000). and Campbell (1996) recorded Tentacuiaria 

There is little known about the tentaculariid coryphaenae Bosc, 1797. Nyhelinia thyrsites 

trypanorhynchs from Australian coastal waters. Korotaeva, 1971 and N. s^iymuv Yamaguti, 1952 

The first collections were made by a French from Australian fishes, while Jones and Beveridge 

scientific expedition under the command of (1998) added N. queenslandensis from a 

Nicolas Baudin between 1801 and 1803 carcharhinid shark as a new species. Palm (1999) 

(Bcvendgc & Campbell 1996), but the ccstodes, recorded Nybelinia lin/iualis (Cuvier. 1817) from 

Tenwcularia coryphaenae Bosc, 1797 and an southern Australian waters and Speare (1999) 



50 



11. W, PAI.M&I. BEVEKtDGF. 



reported Nybelinia sp from sailfisb. hriophorus 
plunpierus (Shaw & Noddcr, 1792) off the coast 
of Queensland. Tints, a total of five tentaculariid 
species has been described from Australia. 

Last and Stevens (1994) summarised 
knowledge of the Australian chondnchthyan 
fauna, the definitive hosts of trypanorhynchan 
ccstodes, as being extremely rich and consisting 
of 206 species, 54% of them endemic. In terms of 
host-specificity and zoogeography of 
itypanorhyiich eesiodes, ihc Australian fauna 
might therefore harbour a high number of 
endemic and locally distributed lentaculariid 
trypanorhynchs, such as was observed within Ihc 
related family Butetrarhynehidac by Beveridge 
(1990). Palm (|9y9) and Palm and Walter (2000) 
suggested that many tentaculanids, for example 
Tenlacularia coryphuenae and Nybtlinin 
lingualis, exhibit a cosmopolitan or circumtropieaJ 
distribution pattern However, records from the 
south-west Pacific and south-east Indian Ocean 
are scarce. For example, the tcntaculariids 
Tentucularm coryphacnoe, Nybelinia afrienna 
and N. scaliodoni were only recently recorded 
Irom Indonesian coaslal waters (Palm 2000). 
Other records which tmghi confirm a wide 
distribution and possibly a low degree of host 
specificity for these trypanorhynchs are still 
lacking. 

The present study was earned out to examine 
ihc species of Nybelinia, Heteronybelina, 
Mixonybelinia and Kotorelta deposited in 
museum collections m Australia as well as in the 
collection of the junior author. Where necessary, 
the specimens are described and illustrated as 
emendations to currently available descriptions. 
Apart from ihc establishment of new host and 
locality records, species identifications provide 
further insight into the zoogeographical 
distribution of lenUculariid eesludes. The 
comparison of the specimens with earlier 
descriptions allows comments to be made on the 
extent of intraspecific morphological variability 
within tentaculariid trypanorhynehs. Together 
with the study of deposited lenlaculariids from 
other collections (Palm 1999: Palm & Overstreet 
2000; Palm and Walter 1999, 2000), the present 
study summarises the current stale of knowledge 
on the genera of tentaculariid cestodes. 



Material Ano Methoo.s 

Specimens were examined from the Australian 
llelm'mthoiogical Collection (AHC) housed in the 



South Australian Museum (SAM), Adelaide, as 
well as the Queensland Museum (QM), Brisbane 
Special attention was given to unidentified 
specimens deposited simply as Nyln-liitiu sp. 
Unmounted specimens were stained in Celestine 
Blue, dehydrated in ethanol. cleared in methyl 
salicylate- and mounted in Canada balsam. Pieces 
of strobilac were embedded in paraffin, seii.il 
sections were cut at a thickness of 5 um and 
stained with haeinaloxylin and eosin, Drawings 
were made using an Olympus BH2 microscope 
wilh an ocular micrometer and drawing lube 
Terminology for morphological features peculiar 
to the Ttypanorhyneha follows LMIfus (1942) and 
Campbell and Beveridge (1994) 

The following measurements were made scolex 
length (SL), scolex width at level of pars 
bolhridialis (SW). pars boihridialis (pbo), pars 
vaginalis (pv), pars bulbosa (pb), pars posibulbosa 
(ppb), velum (vcl), appendix (app), bulb length 
(RL) bulb widlh (BW), bulb ratio (BR), scolex 
proportions of pbo:pv:pb (SP), tentacle width 
(TW) and tentacle sheath widlh (TSW). If 
possible, the tentacle length (TL) was estimated 
The tentacular armature was described as follows: 
armature henneumorphous or hctcromorphous, 
number of hooks per half spiral row (hsr), toi.il 
hook length (L) and the total length nf the base of 
the hooks (B). The abbreviation ; nrn' (not 
measured) indicates lhal no measurement was 
taken All measurements are given in micrometers 
unless otherwise indicated Specimens belonging 
to the same species from different hosts or 
localities were measured in the same order as cited 
in the material examined. If more than three 
measurements were taken from a single host 
species, the mean is given wilh the range in 
parentheses. Because tentaculariid cestodes are 
morphologically uniform and the genera are well 
defined, the species descriptions presented here 
are restricted to key differential features 
Illustrations are provided if useful for future 
.species identification; otherwise the reader is 
leferred to illustrations given by other authors, 
The classification utilised follows Palm (1995. 
1497). Host identity follows Last and Stevens 
(1994) for the elasmobranchs and FishBasc I99S 
(Froese & Pauly 1998) for the teleosts. 



Systematics 

A total of 19 species were identified, and II 
new locality and 20 new host records were 
established. Seven now spcci&s arc described and 



THNTACUI-ARIIDTRYPANORHYNCHS FROM AUSTRALIA 



SI 



a new lenlaculariid genus, is erected: Nybelinia 
hemipristis sp nov., N. mehlhomi sp, nov,, N. 
vicwn'ae sp. nov., Heteronybtlirna auaralis sp. 
nov.. H. pscudorobusta sp. nov.. Mixonybelinia 
cribbi sp. nov. and Kounellwllu jtmesi gen. ct sp. 
nov. Detailed information on individual 
specimens with comments on their taxonomy and 
distribution are provided below. 



Order TRYPANORHYNCHA Dicsing, 1863 

Superfnmily TENTACULARfOlDKA Poche, 
1926 

Family TENTACULARIIDAE Poche, 1926 



Cicnus Nybelinia Poche, 1926 

Nybelinia acquidentata Shipley & llorncll. 1906 
(Pigs 1-2) 

Material examined 

From Dendrociurus zebra (Cuvier, I 829): I 
posllarva, Noumea. New Caledonia, Nov. 1997, 
coll. S. Pichelin (QM G 21803 1). 

Description 

SL=7900 (Pig. 1), SW = 1775; pbo = 2975 
p\' = 4950; pb = 1575; vel = 925, upp = 1-400 
BL=1417 (1400-1450), BW = 280 (270-290) 
BR = 5.1:1; SP= 1.9:3.1:1. Basal tentacular 
swelling absent. TW basal = 100-110, TW 
metabasal = 110-120. Tentacle sheaths straight. 
TSW= 80-100, prebulbar organs and muscular 
rings around basal part of tentacle sheaths not 
visible, Retractor muscles originate in basal parts 
of bulbs. Tentacular armature homeoacanthous, 
liomeomorphous (Fig. 2); distinctive basal 
armature absent. Hooks falciform, with slender 
shaft, stout base and strongly recurved rip, 
L = 57.5-65.0; B= 17.7-22.5. Hooks decrease in 
si/c towards apical region and towards base 
L = 35.0-57.5; B = 12,5-17.5. Smallest hook 
measured L = 1 7 J5; hsr = 7-8. 

Remarks 

The present specimen has one of the largest 
scolcees among Nybelinia species. In addition, the 
posllarva is characterised by falciform hooks over 
60 in length and a bulb ratio greater than 4. The 
posllarva is most similar to Nybelinia 
cuauuleniaru Shipley & Honiell. 1906, described 
by Shipley and llomell (1906), Pintner (1927) and 



Palm (1999). Shipley and Hornell (1906) recorded 
the lype specimen as having a scolex 4-5 mm 
long. The hook shape of the present specimen 
corresponds with those of the type, though the 
hooks in the present specimen are larger (57-65 
compared with 49 in the type according to Shipley 
and Honiell (1906) or 33-38 reported by Palm 
(1999). Pintner (1927) redescribed the type and 
added information on the bulb ratio (4.3:1), hook 
length (48) and scolex measurements (SW = 1900, 
pbo =1500, pb=1600 and vel = 600). He also 
remarked that the hooks were not uniform in size 
hut increased from the base towards the metabasal 
region and then decreased in the apical region of 
the armature. Thus, although the scolex and hook 
sizes of the present specimen are distinctly larger 
than those described from the type specimen, it 
has been identified as N. aequideniata. The 
present finding represents a new host and locality 
record. 

Nybelinia syngenes Pintner, 1929 is the only 
other species of Nybelinia which has been 
described as having tentacular hooks with a length 
greater than 60 and with a similar hook form and 
armature pattern. However, the scolex of this 
species, described originally from Sphvrna 
zygaena (Linnaeus, 1758), is distinctly smaller 
Whether N. acquidentata exhibits variation in 
scolex size, as described for N lingualis (Cuvier, 
1817) and Heteronyhclinia yantaguti (Dollfus, 
1960) (see Dollfus 1942: Palm el a I. 1997; Palm 
1999; Palm & Walter 2000) is not known. Palm 
(1999) has suggested that this might be the case 
for a postlarva from Lepturacanrhus savalu 
(Cuvier, 1829) with a scolex length of 3400. a 
bulb ratio of 3.3:1 and slender falciform hooks 
33-38 in length. 



Nybelinia africana Dollfus. 1960 
(Figs 3-5) 

Material examined 

From unidentified shark: 1 adult, Queensland, 
coll. J. C. Pearson (QM G 218169). 

Description 

SL = 1780 (Fig. 3); SW = 1134; pbo = 930, 
pv = 840; pb = 523: vel = 417; BL = 439 (423- 
459); BW=163 (157-171); BR = 2.7:1. 
SP= 1.81.6.1. TW metabasal = 47-50. Basal 
tentacular swelling absent. Tentacle sheaths 
spirally coiled; TSW = 65-70. Prebulbar organs 
absent, muscular rings around basal part of 
tentacle sheaths not seen. Retractor muscles 



52 



H. W. PALM & 1. BEVERIDGE 







FIGURES 1-2. Nybeliniti aequidcntata Shipley & Hornell, 1906 from Dendmchirus zebrtie. QM G 218031. 1. 
Scolcx. 2. Homeomorphous armature with falciform hooks. Scale bars: Figure 1. 500 um; Figure 2, 100 urn. 
FIGURES 3-5. Nybelinia africana Dollfus, I960 from an unidentified shark. QM G 218169. 3. Scolex. 4. Basal 
and mctabasal armature, 5, Matuie proglottid. Scale bars: Figure 3, 200 um; Figure 4, 20 um; Figure 5, 100 um. 



TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 



S3 



originate in basal parts of bulbs. Tentacular 
armature consists of homeomorphous hooks along 
tentacle (Fig. 4) Tentacles nol completely 
evaginated: metabasal armature differs distinctly 
from basal armature. Basal 2-3 rows ol rosethom- 
shaped hooks with distinct anterior extension of 
base. L= 14-16, 16-17: metahasal hooks larger, 
l.ikitorm with small base, strongly recurved at tip, 
L = 26-27; B = 10-13. Strobila 57 mm long, with 
about 2 1 5 acraspedole segments which enlarge id 
size towards end of sirobila. maximum width 1.48 
mm; velum straignt or very slightly scalloped; first 
segments 15-30 x 600-675. mature segments 
(Fig. 5) wider than long, 220-470 x 650-970 
(n-4). Pre-gravid segments (with some Cggs) 
440-670 x 1 150-1480 (n = 5). terminal pre-gravid 
scL-ment with rounded end. In mature segments, 
genital pore vcnlro-submarginal, in anterior half 
ur anterior third of segment; genital pore to 
anterior end 90-150 in mature and 180-200 in 
pre-gravid segments, genilal pores alternate 
irregularly Cirrus sac thin-walled, elongate, 370- 
480 x 70-100 <n=5), length; width ratio 4.5:1 
(3.7-50:1), distal pole directed ariteromedially. 
nearly reaching antenur end of proglottid, cirrus 
unarmed; internal and external seminal vesicle 
absent. Vas deferens greatly colled, extends to 
midline of segment, then posteriorly towards 
female genital complex. Testes ot varying shape, 
60-90 in diameter, ananged in single layei 
centrally and in double layer peripherally; testis 
number 50-71 per segment, between 5 and 8 
testes anterior to cirrus (n = 5). Ovary follicular, 
in centre of segment, bilobed, each lobe 190-260 
x. J40-220. increasing in size along strobila. 
Vitelline follicles encircle medulla, follicles 30- 
50 in diameter. Ventral and dorsal osmoregulatory 
canals 30 in diameter, testes extend external to 
ventral canal but nol beyond dorsal canal. 

Kcmirks 

The present specimen closely resembles N. 
africana Dollfus, I960 as described by Dolllus 
(I960) and Palm ( 1999) in having (osethoni-shapcd 
basal hooks, changing to falciform metabasal hooks 
bf similar size and shape, Additionally, the segment 
morphology with number and size of segments, 
size of cirrus sac, testes and vitelline follicles, as 
well as arrangement of the genital complex is 
similar. However, some differences were observed 
between the present specimen and A', africana. The 
specimen from Australia differs in having 2 to 3 
rows of basal hooks with a distinct atucrtor 
extension of the base (Fig, 41, which has not been 
reported in other specimens of A', africana 



However, Dollfus (I960, figs 14, 17) illustrated 
comparable basal hooks, some with an anterior 
extension of the base. The seolex is larger in the 
Australian specimen (1780) compared with other 
material (.1118-1568) (Palm et al. 1997) and thr 
testis number is smaller However, as most other 
morphological characters correspond, the specimen 
is identified as N. africana and the observed 
differences are considered to be due to intraspecific 
morphological variability. The present finding is a 
new locality record for the species. 



Nybelinia hemipristis gp. nov. 
(Figs 6-9) 

types 

Holotype from stomach of Hemipristis elongata 
Klunzinger, 1871, Balgal, Queensland, coll. B. Q. 
Robertson, I6.ix.1985 (SAM AHC 28309). 
Paratypes. 3 specimens, Hemipriuit elottgaius. 
Marchinbar Island, Northern Territory, coll. G, 
Cuthbcrtson, 25.V.1985 (SAM AHC 28310). 

Male rial examined 
Typos. 

Description 

SL=2030 (Fig. 6); SW = 1230; pbo= 1000: 
pv = 860; pb=560; ve| = 63il: BL = 553 (550- 
560), BW = 218 (210-220); BR = 2 .5:1; 
SP = 1 8- 1.5:1. Tentacles short, massive, 
decreasing in diameter towards apical region, 
basal tentacular swelling absent. TW basal and 
metabasal = 75-85, TW apical = 37.5-62.5. 
Tentacle sheaths straight (TSW = 50-60), 
prcbulbar organs and muscular rings around basal 
part of tentacle sheaths not seen. Retractor 
muscles originate in basal parts of bulbs. 
Tentacular armature homeoacanthous. 
homeomorphous (Fig. 7); distinctive basal 
armature absent (Fig. 8). Hooks slender, 
rosethom-shaped with anterior elongation of base, 
increasing in size from basal (L= 15 5-30.0; 
B = 163-20 0) towards metabasal rcgion> 
(L = 35.0^0.0; B = 25.0-27.5), then decreasing 
in size towards apical (L = 27.5-32.5; B = 15.0- 
20.0) region; hsr = 6. Strobila acraspedole, velum 
straight; 33 mm long, maximum widlh 1030. 
number of segments 300. Mature segments (Fig. 
9) wider than long, 600-670 x 750-920 Genital 
pore submarginal, 200 from anterior end. 
alternating irregularly. Cirrus sac short and stout. 
180-210 x 50-100 (n = 5). length.width ratio 
2.9:1 (2.0-3.6:1), distal pole directed 



V. 



H. W. PALM &[. REV ER1D0B 




FIGURES 6-9. Nybelinia hemipristii sp. nov. from ffemiprixtis elongata. Holotype, SAM AHC 28309. 6. Scolex. 
7. MclDbasal arrnaturc. 8. Basal armature. 9. Mature pcoglotiid. Note short cirrus sac, the uterine duct (ud), and the 
Mehlis gland (mg>. Scale bars: Figure 6, 200 urn; Figure* 7-8. 40 urn; Figure 9, 100 urn. FIGURES 10-11. 
Nybelinia juxapuulazariahi Reimer, 1980 from a 'sole'. QM G 207318. 10. Metanasal armature. U. Basal 
armature. Scale bars: Figures 10-1 1. 10 urn. 



TENTACULARIIDTRYPANORHYNCHS FROM AUSTRALIA 



55 



anteromcdially, not reaching anterior end of 
segments, cirrus unarmed; internal and external 
seminal vesicle absent. Vas deferens coiled, 
extends to midline of segment, then posteriorly 
towards female genital complex. Testes 84—97 in 
number. 55-85 in diameter centrally and 40-60 
peripherally, distributed in single layer; arranged 
in 2 lateral groups, confluent posterior to ovary; 
extend between ovarian lobes, about 6-7 testes 
anterior to cirrus sac Ovary in centre of segment, 
ovarian lobes 200-260 x 120-160. Vagina ventral 
to cirrus sac. Vitelline follicles encircle medulla; 
follicles 25^40 in diameter. 

Remarks 

The present specimens, witb roscthom-.shaped 
hooks diminishing in size towards the apical 
armature and a strobila consisting of acraspedote 
segments, resemble N. anthicosum Heinz & 
Dailey, 1974. However, the species differ in 
having different scolex forms, tentacles, tentacular 
armatures and sizes, N. anthicosum has an 
elongated prominent velum, long tentacles and 
tentacular hooks which are distinctly spaced (see 
Heinz & Dailey 1974). The largest hooks occur in 
the 7th to 9th rows, N. hemipristis sp. nov. has a 
massive scolex with a shorter velum (Fig. 6), short 
tentacles and tentacular hooks which are tightly 
spaced along the tentacle The largest hooks are in 
the 10th to 14th rows. In the last few hook rows, 
the size and number of hooks per half spiral row 
decrease rapidly (Fig. 7). In N. anthicosum the 
hook size diminishes gradually towards the apical 
part of the tentacle. As with ,V. unthwmum, N. 
httnipristis is a species without a characteristic 
basal armature and witb a smaller basal than 
mctabasal hook size, therefore belonging to 
species subgroup 'An' of Palm ( 1999). 

The description of the strobilar characters of N. 
hemipristis demonstrates that this species hns a 
very characteristic, short, stout cirrus sac. with a 
length: width ratio of 2 in some segments, a feature 
unusual in most species of Nvhelinia. However, 
this character might prove to be of taxonomic 
significance in the future. 

The new species is named after the 
elasmobranch host genus, Hemipristis. 



Nybeliniu jayupauUizuriuhi Reimcr, 19S0 
(Figs. 10-11) 

Material examined 

From 'sole', cither Synaptuta mgru Macleay, 
1880 or Asetaggodes mat leayunu.\ (Ramsay, 



1881): t postlarva, Moreton Bay. Queensland. 
colL J. C Pearson, 1968 (QM G 207318). 

Description 

SL=1060; SW = 700; pbo = 470; pv = 450; 
pb = 365; BL = 323 (320-325); BW=»19 (85- 
135); BR = 2.7:1; SP= 1.3:1.2.1. TW = 32.5-35; 
hsr=6. Basal tentacular swelling absenL Retractor 
muscles originate in basal parts of bulbs. Tentacular 
armature consists of humeomorphous slender 
uncinate hooks, L= 15.0-16.3. B = 1 1 2-13.7 (Fig. 
10). Size of hooks diminishes towards basal part of 
tentacle, L = 1 1.2-13.7. B = 8.7-1 1 .2 (Fig 1 1 ). 

Remarks 

N. jayapaulazariahi was originally described 
from Cynnglossus sp. from the Bay of Bengal, 
India by Rcimer (1980). Palm (1999) redescribed 
the species from another host, Harpodon nehereus 
(Hamilton-Buchanan, 1822), also from India. The 
tentacular armature has characteristic, slender, 
regularly curved hooks which increase in size 
towards the metabasal part of the tentacle. The 
present specimen, also from a 'sole', has a similar 
hook shape (compare Fig. 4 with Fig. 5 of Palm 
1999) and bulb ratio, while it differs in having a 
larger scolex (1060 in the present specimen 
compared with 530 described previously) and 
hook size (11.2-16.3 in the current specimen 
compared with 5.6-1 1.2 in previous descriptions). 
However, the present specimen was in a poor state 
of preservation, and the larger scolex might be 
related to the slightly larger hook size. A small 
scolex of about 0.5-1.0 mm, together with the 
uncinate hook form, is characteristic for N. 
jayapaulazariahi, and soles seem to be important 
intermediate hosts. The present finding represents 
a new locality record for the species, 



Nyhelinia mehlhomi sp. nov. 

(Figs 12-14) 

Types 

Holotype from stomach of Hemigaleus 
microstoma Bleeker, 1852. Heron Island, 
Queensland, coll. P. McBoarnidn, 20.xii.1995 
(QM G 218032); paratype, same dala (QM G 
218033). 

Material examined 
Types. 

Description 
SL = 560 (Fig. 12); SW = 400; pbo = 330; 



56 



H. W. PALM & I. BEVERIDGE 




12 





FIGURES 12-14. Nybelinia mehlhorni sp. nov. from Hemigaleus microstoma. Types, QM G 218032-3. 12. 
Scolex. 13. Metabasal and apical armature. 14. Mature proglottid. Note the uterine duct (ud) and the seminal 
receptacle (sr). Scale bars: Figure 12, 50 um; Figure 13, 10 um; Figure 14, 50 urn. FIGURES 15-16. Nybelinia 
scoliodoni (Vijayalakshmi, Vijayalakshmi & Gangadharam, 1996) from Diodon hystrix. QM G 218035-7. 15. 
Scolex. 16. Basal and metabasal armature, external surface. Scale bars: Figure 15, 150 um; Figure 16, 25 um. 



TENTACULAR! ID TRYPANORHYNCHS FROM AUSTRALIA 



f? 



pv=125. ph = 250; vcl = 200; BL= 178 (150- 
200); BW = 58 (50-70); BR = 3.1.1; 
SP = 1.3:0,5:1. Tentacles short; basal tentacular 
swelling absent; TW = 22.5-27.5. Tentacle 
sheaths sinuous to spirally coiled (TSW = 17.5- 
20.0); muscular rings around basal part of tentacle 
sheaths seen. Retractor muscles originate in basal 
parts of bulbs. Tciuacular armature 
homeoacanlhou.s, homcomorphous (Fig. 13), 
distinctive basal armature absent. Hooks 
falciform, with stout base, elongated handle and 
strongly recurved tip. Hooks increase in sue from 
basal lk = 3.5-7.5. B = 2.5-5.0) to metabasal 
region (L = 15 0-17,5, B =6.3-8.7) then decrease 
towards- apex [t- II. 3-13.8, B = 2.5-4. 3); 
hsr - 7. Small worms, length 25 (16) mm, 
maximum width 950 (700) with 135 (90) 
segments. Strobila acraspedote, velum straight; 
pje mature (testes visible) proglottid* (Fig. 14) 
80-280 long x 560-790 wide; mature segments 
(female genitalia developed) wider than long, 
380-600 x 600-900; terminal segment 830 x 800. 
Genital pore almost lateral, 1 30-200 from anterior 
end (190 m terminal segment); pores alternate 
irregularly. Cirrus sac elongate, 170-220 x 40-60 
(n = 5) in mature segments. Iength:width ratio 
4.1:1 (2.8-5.5:1)," distal pole directed 
aitictotnedially, not reaching anterior end of 
segments; cirrus unarmed, internal and external 
seminal vesicles absent. Vas deferens greatly 
onli-.d in midline of segments, extending 
posteriorly towards female genital complex. 
Testes 103-120 in number, 80 x 95 in Sue, 
smaller testes peripheral, 35-40 in size. 
distributed in single layer; testes in 2 lateral 
groups confluent posterior to ovary; 10-16 testes 
anterior to cirrus sac. Vagina tubiform. 30 in 
diameter, ventral to cLrrus sac. passes 
aiiicromedially to midline, then posteriorly to 
ovary, seminal receptacle present, 60-90 x 25-50. 
Ovary hi Centra of segment, ovarian lobes 250- 
300 X 160-210. Vitelline follicles encircle 
medulla, follicles 30-45 in diameter, Uterine pore 
absent 

Remarks 

Following Palm (1999), N. mehlhorm sp. nov. 
belongs 10 Nxbelima species without a 
characteristic basal armamre and a basal hook size 
smaller than the metabasal one, thus belonging to 
the subgroup 'Aa'. 

On the basis of the hook shape, the species 
lescmbles N. aequidenlala (Shipley & Hornell, 
1906) and N. gareensis Dollfus, 1960. While the 
former is much larger than the present specimen, 



the latter species has different scolex proportions 
as well as a different hook arrangement, as 
illustrated recently by Palm and Walter (2000) 

A*, mehlhomi sp. nov, also resembles N, 
berigalensis Rcimer. 1980 in its falciform hooks 
and in hook size (Reimcr 1980). However, A' 
benfialen'ii.y can be distinguished by having a 
distinctly different scolex form and size, a bulb 
ratio of about 2. long and slender tentacle sheaths 
and tentacles with spaced falciform hooks (Rcimer 
1980). By contrast. N. mehlhorm sp. nov. has 
short tentacles with hooks tightly spaced along the 
tentacle. The scolex (0.56 mm in total length) is 
much smaller. 

The new species is named in honour of Prof. 
Heinz Mchlhorn. Hcinricli-Heinc-Univcrsicv. 
DOsseldorf. 



Nyhetinia schmidti Palm. 1999 

Material examined 

From Isurus oxyrhinchus Ratmesquc, 1810; 1 
adult, Bicheno, Tasmania, coll. B. G Robenson. 
24.iv.l987 (SAM AHC 28313). 

Description 

SL = 1600; SW = 850; pbo = 1040. pv = 680; 
pb = 340; vcl = 600; BL = 298 (270-330): 
BW= 131 (110-150): BR = 2.3:1; SP= 3.0:2.0:!. 
TW basal =45-48, TW metabasal = 30.0-32.5: 
hsr = 5-6. Basal tentacular swelling absent 
Tentacle sheaths spirally coiled, TSW = 25-40 
Prebulbar organs absent, muscular rings prcsehi 
around the tentacle sheaths. Retractor muscles 
originate in basal part of bulbs. Tentacular 
armature homeoacanthous, homcomorphous. 
consisting of massive rosethom-shaped hooks 
with anterior extension of base, size in metabasal 
region L= L3.8-15.0, B= 11.3-12.5. decreasing 
towards hase, L = 8.8-1 1.3, B = 6.2-8.8. 
Characteristic basal armature absent. Strobila 
immature with few segments, acraspedote: 
segments Wider than long (660-690 x 15-60). 

Remarks 

The present specimen closely resembles that 
of the type, also from Isurus uxyrhinchus (syit. 
Isurus glaucus), in having similar scolex 
proportions and almost identical hook sizes. 
Other species with a similar tentacular armature 
are Nybelinia strongyla Dollfus, I960 and 
Heteronybelinia oust talis sp. nov. {vide infra), 
While the latter species differs in having 
homcomorphous hooks of different sues on the 



S8 



H. W. PALM & I BEVER1DCH 



internal and external tentacle surfaces as well as 
smaller basal hooks, N. strongyla differs in 
scolex proportions and larger hook sizes. The 
present record extends the range of distribution 
from the South African to the southern 
Australian coast. 



Nybelinia scoliodoni ( Vijayaiakshmi, 
Vijayalakshnii & Gangadharam, 1906) 
(Figs 15-16) 

Material examined 

From Diodon hystrix Linnaeus, 1758: Heron 
Island, Queensland, 2 postlarvae, coll J. Sakanari, 
23.viii 1986 (QM G 218034): 8 postlarvae, coll. 
M. K. Tones, 6 vii.1995 (QM G 218035-218037); 
8 postlarvae, coll. T. 11 Cnbb, Tan. 1997 (QM G 
218038-218041). 

From Makaira tndica (Cuvicr, 1832): I 
postlarva. Cape Bowling Green. Queensland, coll 
P. Speare, 18.ix.1987 (SAM AHC 21351). 

From Istiophoras plunpierus (Shaw & Nodder, 
1792); Whitsunday Island, Queensland, coll. P. 
Speare. 5xii.l988 (SAM AHC 21351). 

From Carcharhinus linibatus (Valenciennes, 
1839). I adult, stomach. Bremer Island. Northern 
Territory, coll. J Stevens, 29.V.I985 (SAM AHC 
28314), 

Description 

SL=1350, 1115 (Fig. 15); SW - 860, 800; 
pbo = 670, 520; pv = 230, 210, pb = 300, 320; 
vcl = 330, 330; app=3l0, 200; BL = 288 (280- 
300), 282 (260-300); BW= 115 (110-120), 125 
(120-130); BR = 2.5: 1. 2.3:1; SP - 2.2.0.8:1 . 
1 .6:0. 7:1. Tentacles nearly completely evaginaicd, 
TL = 390. 375; basal tentacular swelling absent. 
TW at basal armature 35, 30; TW at mctabasal 
armature 50, 45; TW at apex 30, 25. Tentacle 
sheaths sinuous, TSW = 30, 40; prebulbar organs 
and muscular rings around basal part of tentacle 
sheaths not visible. Retractor muscles originate in 
basal parts of bulbs. Mctabasal armature 
homcoacanthous, homeomorphous; distinctive 
basal armature present (Fig. 16). Basal armature 
consists of about 10 rows with compact rosethom- 
shaped hooks, increasing in size from L = 4.i, 
B = 3.9 (row I) to L= 19.5. B = 1 1.7 (row 10). 
From row 11, hook form changes to slender 
spiniform, L = 22.7. B-13.0. Meiabasal and 
apical hooks L=29.9. with smali base, B- 11. 
Number of hooks per half row (hsr) 6-7 in basal 
region, decreasing to 5 in mctabasal and apical 
regions. 



Remarks 

Nybelinia scoliodoni is a widely distributed 
tentaculariid and is here recorded for the first time 
from Australian waters. The species is easily 
recognised by its rosethom-shaped basal armature 
of about 1 1 rows which is followed by long, 
spiniform metabasa) hooks. The present 
specimens correspond with the description given 
by Palm (1999) in having a similar armature and 
bulb ratio. They differ in having a larger scolex 
and hook size. However, they correspond closely 
to the original description of Nybelinia 
(= Tentacularia) scoliodoni of Vijayaiakshmi ct 
al. (1996). The present findings constitute three 
new host records. C. hmbutus has been recorded 
previously as a definitive host for N. scoliodoni 
by Palm (1 999). The specimens described here are 
the same as those reported by Speare (1999). 



Nybelinia strongyla Dollfus. 1960 
(Figs 17-20) 

Material examined 

From Argyrosomus hololepidutus Lacepcdc, 
1802; 2 postlarvae from stomach, Murray Mouth, 
South Australia, coll. M. G O'Callaghan. May 
1992 (SAM AHC 28345). 

From Jolimus vogleri (Blecker, 1853): 1 
postlarva from body cavity. Heron Island, 
Queensland, coll J. Reddich, Jan 1997 (QM G 
218109). 

Description 

Postlarvae from A. holoUpidotus: SL = 1303, 
1566 (Fig. 17); SW = 648. 796; pbo = 700. 781; 
pv = 647, 764; pb=292, 355; vel - 332. 326; 
app = 374, 429; BL = 286 (244-292), 355 (334- 
380); BW= 86 (82-90). 98 (92-107); BR= 3.3 1. 
3.6 1; SP = 2.4:22:I, 2.22 2:1, Tentacles not 
completely evaginated; basal tentacle swelling 
absent. TW =50-58, 43-45. Tentacle sheaths 
sinuous, TSW = 36^10, 36-40; prebulbar organs 
and muscular rings around basal part of tentacle 
sheaths not visible. Retractor muscles originate in 
basal part of bulbs. Tentacle armature 
homeoacanthous, homeomorphous (Fig. 18); 
distinctive basal armature absent. Hooks slender, 
rosctliorn-shaped, increasing in size from basal 
towards meiabasal part of tentacle. Metabasal 
hooks L= 17.5-20.0: B= 13.8-16 3; basal honks 
L= 10.0-12 5; B= 10.0-12.5; hsr = 7 

Postlarva from ./. voglen: SL=IKK0; 
SW = 1660; pv = 680; pb = 440; BL = 308 (300- 
325): BW-129 (115-150); BR = 2.4:1; 



TENTACULARIIDTRYPANORHYNCHS FROM AUSTRALIA 



59 




FIGURES 17-18. Nybelinia slrongyla Dollfus, 1960 from Argyroaomux hohlepidotus. SAM AHC 28J545. 17 
Scolcx. 18. Metabasal armature. Scale bars: Figure 17, 150 um; Figure 18: 15 urn. FIGURES 19-20. Nybelinia 
mrangylu from Jnhninps vogleri. QM G218109. 19. Melabasal armature. 20. Basal armature. Scale bars: Figure 
19-20: 10 urn. FIGURES 21-24. Nybelinia thyrxites Korotacva, 1971 from Trachurus dedivis. QM G 214194-5. 
21. Scolcx. 22. Basal armature. 23. Metabasal armature with falciform hooks. 24. Apical armature with rosethorn- 
shaped hooks. Scale bars: Figure 21, 200 urn; Figures 22-24, 25 urn. 



60 



H. W PALM & I HRVERIDCE 



SP = -:l.5:l. TW basal = 37-40, TW 
mctabasal = 30-33. Basal tentacular swelling 
ahscnt. Tentacle sheaths straight; TSW = 20-25. 
Prebulbar organs absent, muscular rings around 
basal part of tentacle sheaths absent. Retractor 
muscles originate in basal part of bulbs, Armature 
homeoacamlious, homeomorphous (Fig 19); 
characteristic basal armature absent (Fig 20). 
Honks rosethorn-shaped with anterior extension 
of basal plate, hooks in basal part of tentacle 
smaller (L = 7.5-10 0; B = 7.5-9.5) than in 
mctabasal (L = 13.8-16.3, B= 10.0-12.5) 
armature. Number of hooks per half spiral (hsr) 
diminishes from 6-7 in mctabasal region to 5-6 
in apical part of tentacle. 

Remarks 

Nyheliniu sirongyla Dollfus, 1960 was 
described from a postlarva from the wesl coasl of 
Africa, and is characterised by the 
homeoacanlhous tentacular armature with slender 
rosethom-shaped hooks, reaching a size of up to 
16 ifi the mctabasal armature, Palm and Walter 
(20001 described aduli .specimens of N. sirongyla 
from Africa. The present specimens correspond 
closely in their tentacular armature as well as in 
scolex proportions to those specimens described 
by Dollfus (1960) and Palm and Walter (2000) 
Minor variations in the hook pattern are observed 
within N. strong\labal are difficult to define, and 
many scolex characters within tentaeulariids 
appear to be variable (Palm 1999. Palm and 
Walter 2000). Therefore, the posllarvae described 
above are identified as N. sirongyla. However, 
some doubt surrounding the identification 
remains. Further studies are needed to determine 
whether posllarvae with rosclhom-shapcd hooks 
as described above are conspecilic with the 
African material, or whether they belong to a new 
species of h'ylwlinia. Ar^yrosanws hololepidotus 
is a new host record, and the distribution of the 
species is extended to the southern Australian 
coast. 

The specimen from J. vogleri, with roseihorti- 
shaped hooks, resembles N. iingnalis, N. schmidti. 
N. sphxmac. N. strongyla and N. thyrsiies. It 
differs from A', Hngualis in having basal hooks 
with an anterior extension of Ihe basal plate, and 
from /V ihxrsiws in having differently shaped 
nietabasal hooks and a larger bulb ratio (sec 
below). N. schmidti differs io having smaller 
hooks and different scolex proportions. N. 
sphytnae differs in having slender, more uncinate 
hooks in the mctabasal armature, while the hooks 
arc more massive rosethorn-shaped in N. 



sirongyla. In the present specimen, the basal hook 
length is slightly smaller than described by Palm 
and Walter (2000) for N. sirongyla. However the 
metaoasal iiouks are Ihe same sue Although the 
scolex proportions were difficult to measure in 
the present specimen, il is also identified as N 
strongyla, representing a new host and locality 
record 



Nybelinia thyrsites Korotaeva, 1971 
(Figs 21-24)' 

Material examined 

From Trachurus deelh'is Jenyns, 1841: 2 
postlarvae from stomach wall, Crayfish Bay, 
Tasmania, coll. K. B. Sewcll, ll.\1987 (QM G 
214194.212145). 

From Carcharbinin braihyurus (Gutither, 
1870): 1 adult from stomach. Goolwa, South 
Australia, coll R. Martin, 2S.X.1985 (SAM AHC 
28312). 

From Mustcluf, aniaalicus Gtinther, 1870: 1 
adult from siomach, Goolwa, South Australia, 
coll. R. Martin, 28.x. 1 985 (SAM AHC 283 1 1 ). 

Description 

SL=2032. 1550 (Fig. 21); SW=l520, 700: 
pbo= 1024. 610; pv = 640, 500; ph = 368. 53ft 
veJ = 992, 490; app = 496; BL = 358 (355-365). 
495 (490-500): BW = 1 17 (1 14-120), 155 (140- 
170). BR = 3 11, 3,2:1. SP = 28I 7:1, 1 .2:1:1. 
Tentacle sheaths spirally coiled; TSW =51-57, 
35-45. Prebulbar organs ahseni, muscular rings 
•around basal part of tentacle sheaths present. 
Reuactor muscles originate in basal part of bulbs. 
Tentacles long (TL = 832) and slender, basal 
tentacular swelling absent; TW basal = 38-44, 
25-30: TW nietabasal = 57.0, 57.7-62.5; TW 
apical 32.0, 27-5. Chaiacteristic basal armature 
present (Fig 22), consisting of about 7-8 rows of 
homeomorphous, rosethorn-shaped hooks with 
slight anterior extension, increasing in size 
towards nietabasal armature (1st row hooks: 
L = 9.5-1 1.0, B = 8 0-1 1.0, L= 7.5-8.7, B = 0.3- 
7.5; 8th row: L =158-17.4, L= 16 2-17 5. 
B =11.0-12.6, B = 10.0-12.5). Meiahasal 
armature of about 12-14 rows of hooks, with 
largest hooks in rows 12-17. Metabasal armature 
with slender falcate to falciform hooks (L = 21.7- 
26.9 B = 8.7-1 10; L = 21.3-24.0, B=11.3- 
12.5) (Fig 23). Apical hooks rosethorn-shaped, 
with hook form changing from slender rosethorn- 
shaped with slight anterior extension to rosethorn- 
shaped wilh distinct anterior extension (Fig. 24). 



TENTACULARfID TRYPANORHYNCHS FROM AUSTRALIA 



6! 



Hooks diminish in size Towards apical region 
(34lh row of hooks; L= 14.2-15.8. B = 11.0- 
14.2); hsr hasal: h-7; hsr metabasal and apical: 6. 

Remarks 

N. thyrsites was redescribed in detail by 
Bcveridge and Campbell (1996). The presenl 
material corresponds with the redescription given. 
In addition, the hasal hook form differs from ihe 
metabasal and apical hooks; thus, the species 
belongs to subgroup 'Ba' of Palm (1999). The 
present specimens from Trachurus declivis 
provide a further example in which ihe postlarvac 
can be larger than adult worms, and also show 
variable scolex measurements depending on the 
state of contraction. Trachurus declivis and 
Carcharhinus brachyurus arc new hosts for N. 
thyrsites. 



Ns'betinia vicloriue sp. nov 

(Figs 25-26) 

Types 

llolotypc from body cavity of Lepidoirigla 
modesta Waiie, 1899, Port Phillip Bay, Victoria, 
coll. R. Norman, 17.W.1989 (SAM AHC 28343); 
paratypc, same data (SAM AHC 28344). 

Material exam\md 
Types. 

description 

SL = 1030, 1050 (Fig 25); SW=700, 650; 
Pbo = 570, 450; Pv=350, 320; Pb = 315. 320; 
Rl.. = 3IO (300-315), 310 (305-320); BW=124 
(115-135), 116 (1 10-125); BR = 2.5; I, 2 7:1; 
SP= 1,8:1.1:1, 1.4:1.0:1. Tentacles elongate; 
hasal tentacular swelling absent. TW = 40.0-50.0, 
45,0-50.0. Tentacle sheaths sinuous lo spirally 
coiled (TSW = 40.0-50.0, 35 0-40.0); prebulbar 
organs and muscular rings around basal pan of 
tentacle sheaths not seen, Retractor muscles 
originate in basal part of bulbs. Armature 
homeoaeanlhous. homeomorplious (Fig. 26); 
distinctive basal armature absent. Hooks 
falciform, with stout base, elongated handle and 
strongly recurved tip. Hooks increase in sire from 
basal (L = 7.5-10.0, B = 6 3-8.8) to metabasal 
region (L~ 12.5-15 0, B = 7.0-8.0) and decrease 
towards apical (L= 11.3-12.5, B = 6.3-8.0) pan 
of tentacle; hsr = 7. 

Remarks 
This species, with a tentacular armature 



consisting of slender falciform hooks, increasing 
in size towards the metabasal region and 
decreasing in size towards the apical region, 
resembles Nybelinia aequidentata Shipley & 
Hornell, 1906; N. syngews Pintner, 1927; N. 
anantaramanorum Reimer, 1980; and N. 
bengalensis Reimer, 1980. While the former two 
species differ in having different scolex 
proportions and distinctly larger hooks (see 
remarks on N. aequidentata), the presenl 
specimens are most similar to N. bengalensis and 
N. anantaramanorum. N. anantaramanorum has 
larger hooks and was considered a species 
inqtiirenda by Palm and Walter (2000). ft. 
bengalensis differs in having different scolex 
proportions, including very short bulbs (BR<2) 
(Reimer 1980). Thus, the present specimens 
represent a new species here named Nybelinia 
Victoria? sp. nov., the specific epithet being 
derived from the Australian state from which 
samples were collected. With hook size smaller in 
the busal rather than the metabasal region, the 
species is considered to belong to subgroup Aa' 
of Palm (1999). 



Genus Heteronybetinia Palm , 1 999 

Heterunybelinia auslralis sp. nov. 
(Figs 27-30) 

Types 

Holotype from stomach of Carcharhinus 
amboinensis (Mullcr & Henle. 1839), St 
Lawrence. Queensland, coll. B. O. Robertson. 
29.*. 1985 (SAM AHC 28315); paratype, same 
data (SAM AHC 283 1 6) 

Material examined 

From Carcharhinus amboinensis: types. 

From Carcharhinus brachyurus: 1 specimen, 
Coolwa, South Australia, coll. R. Martin, 
28.x, 1 985 (SAM AHC 28317). 

Description 

SL=1190, 1210 (Fig. 27); SW=660. 680: 
pbo=650, 700; pv=530, 650; pb = 350. 310; 
vel=220, 190; BL=313 (30S-320), 308 (290- 
340); BW=138 (125-150), 119 (110-130); 
BR = 2.3:1, 2.6:1; SP = 1.9:1.7:1, 2.3.2.1:1. 
Tentacle sheaths spirally coiled. TSW = 27.5- 
32.5, 25.0-35.0. Tentacles robust, TL = 255-290, 
increasing in width towards apex. TW basal and 
metabasal 30.0-32.5; basal tentacular swelling 
absent Prebulbar organs absent; muscular rings 



62 



H. W. PALM & I. BEVERIDGE 




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FIGURES 25-26. Nybelmia vicwriue sp. nov. from Lejridorrigla rrwdcsia Types, SAM AHC 28343-4, 25. Seolex. 
26. Basal and mctabasal urmauire. Scale bars: Figure 25, 100 urn; Figure 26. 20 urn. FIGURES 27-30. 
Ileteronybeiinia QUSlralis sp. nov. from Carcharhinits ainhninensis , Types, SAM AHC 283 1 5—6. 27. ScolcX, 28. 
Mctabasal armature. V). Basal armature. 30. Mature proglottid Note the uterine pore. Scale bars: Figure 27. 100 
(iin; Figures 28-29. 10 |ilrt; Figure 30, 50 um. 



TENTACULARIIDTRYPANORHYNCHS FROM AUSTRALIA 



rt 



around basal parts ot tentacle sheaths present. 
Retractor muscles originate at base of bulbs. 
Tentacular armature homeoacanlhous. 
heteromorphous, with hooks tightly spaced along 
tentacle (Fig 28); characteristic basal armature 
absent (Fig 29). Hooks massive, rose-thorn 
shaped, with slight anterior extension of base, 
increase in size from basal towards ntetabasal 
armature. Hook size differs on bothridial/ 
mitibothridial (L = 12.5-15.0, B = 11.3-13.7) and 
antihollitidial/hoihridial (L= 1 1.3-12.5, B=7.0- 
8.7) tentacle surfaces (Fig. 28). with two tentacles 
having largest hooks on bothridial surfaces and 
two othei tentacles having largest hooks on 
aiiuhotJuidial surfaces. BasaJ hnoks L = 5.0-7.5, 
7.5-87; b =8.7-10.0 (internal) and L=5.0-6.3, 
5.0-7.5; B = 3.8-5.0, 5.0-6.3 (external); hsr=6- 
7 Strobila only very slightly ( raspedote, velum 
scalloped; total length 18.0 and 15.0 mm, 
maximum width 640 and 730, with 125 and 155 
scgmcnls, In hololype. Icrminsd segment pre- 
mature (320 x 550), segments wider than long, 
231 x 630. Genital pores ventro-submarginal 60 
from anterior margin; alternate irregularly. Cirrus 
sac of pre-mature segments elongate and slender, 
2 10-230 x 35-50 (n = 5), lenglli:width ratio 5.3: 1 
(4.6-6.0:1), with distal pole directed 
anteromedially, reaching anterior end of segment 
(Fig 30); cirrus unarmed; internal and external 
seminal vesicles absent. Vas deferens coiled, 
extends posteriorly from cirrus sac towards female 
genital complex. Tesles 106-130 in number, 
smallest testes at margin of medulla, distributed in 
I -2 layers, in 2 lateral groups, confluent posterior 
to ovary; extend between ovarian lobes; 8-10 
testes anterior to cirrus sac. Ventral 
osmuregulutory canal 15 in diameter, internal to 
sinuous dorsal canal, 5 in diameter; 24 prominent 
bands of longitudinal muscles on each side of 
strobila Segments in paratype mature, wider than 
long (240 x 700 to 380 x 720). Genital pores 1 10 
from anterior margin. Cirrus sac of mature 
segments (Fig. 30) elongate and slender, 260-320 
x 50-«0 (n = 5), length. width ratio 5. 21 (4.3- 
5.8 1). Central testes 15-65 in diameter, smallest 
testes peripheral, 30-50 in diameter, distributed in 
1-2 layers, ovarian lobes 110-200 x 80-110, 
seminal receptacle 50-65 x 25-30. 

Remarks 

Hetemnybelinia ausinilis sp. noV. has a unique 
armature. The tentacular, hooks are tightly spaced 
and appear humeomorphous along the tentacle. 
However, the hook sid:cs differ On the bothridial 
and antibothridial tentacular surfaces, fn addition, 



two tentacles have the largest hooks on the 
bothridial surfaces while the other two tentacles 
have the largest hooks on the antibothridial 
surfaces. Whether this armature pattern is unique 
or whether it occurs in other tentaculariid specie^ 
remains to be determined. 

The specific epithet indicates the occurrence of 
the species in Australian waters 



Ileteronybelmia estigmena (Dollfus. 1960) 
(Figs 31-36) 

Material examined 

From Carcharhinux timbalus Valenciennes, 
1839: 16 adults from spiral valve, GcTaldton, 
Western Australia, coll. B. G. Robertson, 
27.xi.1986 (SAM AHC 21352. 28318); 4 adults 
from spiral valve, Nickol Bay, Western Australia, 
coll. B. G. Robertson, II.xi.1986 (SAM AHC 
28319). 3 adults from spiral valve, Darwin 
Harbour, Northern Territory, coll. B. G. 
Robertson, 28.viii 1986 (SAM AHC 28320); 6 
adults from spiral valve. Fog Bay. Northern 
Territory, coll. B G. Robertson, 4.x. 1986 (SAM 
AHC 28329). 

From Carchurhmus a.mblyrhynclwidcx 
(Whitley, 1934): 2 adults from spiral valve. 
Tommy Cut Mouth, Northern Territory, coll. B. 
G. Robertson, 10 ix. 1986 (SAM AHC 28321). 

From Carcharhiiius sp - 3 adults from spiral 
valve, Queensland, coll. B. G. Robertson. Oct 
1985 (SAM AHC 18322). 

From Sarda auslralis (Macleay. 1881): 9 
postlarvae from gill arches. Heron Island, 
Queensland, coll. M. K. Jones, Jan. 1991 (QM G 
218O42-2I8046). 

Description 

AdttU: SL= 1650, 1500 (Fig. 31); SW = 790, 
770; pho=820, 820; pv=690, 610; pb=4SQ. 
430, ppb=110, 80; vel = 330, 320; BL = 455 
(440-480), 425 (420-430); BW=133 (130- 
140), 120 (118-122); BR = 3.4:1. 3.6:1; 
SP= 1.7:1.4:1. 1.9.1.4:1. Tentacles long, slender 
(TL = 520, 560), TS W = 42.5-47.5. 45.0-55.0: 
TW basal 45-50, 42.5-45; TW apicaJ 25-30. 
27.5-32,5; basal tentacular swelling absent 
Prehulbar organs absenl, muscular rings around 
basal part of tentacle sheaths visible in one 
specimen Retractor muscles originate at bases 
of bulbs. Tentacular armature homeoacanthous, 
hetcromorphous, no characteristic basal armature 
present (Figs 32-33). Hooks roselhorn-shaped. 
increase in size towards metabasal part of 



M 



H. W. PALM & I. BEVERIDGE 




32 



34 










%*& 



' 



w 



6 ( 



FIGURES 31-34. Htteronybelmia esiigmena (Dollfus. I%0; from Carcharhima limbatus. SAM AHC 28318 20. 
31. Scolex 32. Metabasal ;umaiure. 33. Basal armature. 34. Mature proglottid Scale bars: Figure 31, 150 pm; 
Figures 32-33. 15 urn; Figure 34, 100 um. FIGURES 35-36. H, estigmena lrom Sardu australix. QM G 218042- 
46. 35. Scolex. 36. Basal armature. Scale bars: Figure 35. 150 um; Figure 36, 15 um. 



TENTACULARUDTRYPANORHYNCHS FROM AUSTRALIA 



65 



tentacle. Hook form varies from compact and 
rosethorn-shaped (bothridial) lo smaller 
rosethom-shaped hooks with elongated base 
(antibothridial). Hook size, metabasal, L = 1 1,3- 
12 5; B= 9.3-10.8 (bothridial) and L = 9.3-10.8; 
B = 10 (anliboihridial), with largest hooks 
L= 13.8-15 0; B = 10.0-12.5 (bothridial); hooks 
diminish in size towards apical part of tentacle. 
Hook size in basal region L- 7.5—8.8; B=6.3- 
10.0 (hotJiridial) and L = 5.0-7.5; B= 6.3-7.5 
(antibothridial); hsr - 6-7. Strobiia 56 mm long, 
maximum width 1250. about 280 acraspedote 
segments (Fig. 34). First segments behind velum 
short, enlarge in size, mature segments wider 
than long, from 500 x 840 to 700 x 1030. In 
mature segments, genital atrium ventro- 
submarginal, in anterior third of segment; genital 
pon:s alternate irregularly. Cirrus sac elongated. 
30<)_450 x 811-110 (ii = 5), length:widlh ratio 
3.8 4.1.1. directed anitromedialjy, sac thin- 
walled; cirrus unarmed and coiled within sac. 
Ovary median, follicular, ovarian lubes 200-2?0 
(Icmjlh] \ 1 10-170 (width), Mchlis 1 glajid 80 in 
diameter, seminal receptacle 110 x 70; testes of 
variable shape. 60-80 x 30-60 (n a 5) in 
diameter, arranged in single layer; testes number 
106-126 (n = 5) per segment, encircle female 
genital complex and occupy entire medulla 
except lor region ot female genital complex and 
anterior ro it; 7-9 testes anterioT to cirrus sac. 
Vitelline follicles 20-40 in diameter, egg 
diameter 20-25: ventral osmoregulatory canals 
•along margins of cortex. 6 in diameter. 

Posrlana SL=I650. 1500 (Fig. 35). 
SW=790, 770; pho = 620, 820; pv = 690, 610; 
pb = 480. 430; ppb= 110, 80; vel = 330, 320; 
BL=460 (430-470), 445 (440-450); BW=11S 
(110-120), 120 (118-122); BR = 4.0, 3.7; 
SP= 1 7:1 4:1, 1.9:1.4-1. Tentacles long, slender 
(TL = 550-580), TSW = 40-45; TW basal = 35- 
40, TW metabasal and apical = 30-40; basal 
tentacular swelling absent. Prehulbar organs 
absent, muscular rings around basal part of 
tentacle sheaths present. Retractor muscles 
originate at base of bulbs. Tentacular armature 
homeoacanthous, hctcromorphous; characteristic 
basal armature absent Hook shape vanes (mm 
compact, roselhorn-sluiped (bothridial) to smaller 
roscihorn-shaped hooks with elongated base 
(antibothridial). hooks increase in size towards 
metabasal part of tentacle on bothridial and 
antibothridial surface (Fig. 36). Hook size, 
metabasal. L= 125-150. B =11.8-13.3 
(bothridial) and L = 11.3-12,5; B = 7.5-10.0 
Unlibolhndial): basal L = 7.5-10.0; B = 8.8-11.3 



(bothridial) and I. = 5.0-7.5; B = 6.3-8.X 
(antibothridial); hsr=6-7. 

Kenuirkx 

Hetcronybchnia cstigmena (Dollfus, I960) is a 
well-described tentaculariid. Palm (1995) reported 
this cestode as Nybelinia alloiotica Dollfus, 1960 
from Carcfiurlunus limbatus in the Atlantic, the 
same host species as in the present study. Palm 
(1 , ' , '9) and Palm and Waller (2000) provided 
additional information on tbc species. H. 
esii^meiia resembles H. peruleraeus (Shipley & 
HornelU 1906) and H efangtaa (Shah & Bilquees, 
1979) known from the coast of India. However, 
the hook size increases on both (he bothridial and 
antibothridial tentacle surfaces in //. estigmena. In 
H. elottgata, the hook size increases on the 
antibothridial tentacle surface only (Palm A 
Waller 1999). while in H- perideraeus, the basal 
and metabasal hook sizes arc the same (Palm 
1999) The present study represents the first record 
of the cestode from Australian waters. 
Carcbarhinus amblyrhynchoides and Sctrdti 
ausiralis represent new hosts for the species. 



Heleronybelinia pscudorobmta sp. nov. 
(Figs37'-4I) 

Types 

Holotype from gills of Lepidomgla modesta 
Waite. 1899. Port Phillip Bav. Victoria, coll. R. 
Norman, 17.iii. 1989 (SAM AHC 28341); 3 
parity pes, same daw (SAM AHC 28342) 

Mali-rial examined 
Types. 

Description 

Small form (holotype and one paratype): 
SL = 960 (Fig. 37), 730, SW = 660, 550; 
pbo = S70, 450; pv = 420, 270: pb=285, 290; 
app = 220. 160; vel = 70. 60; BL = 280 (275- 
285). 298 (275-315); BW=150 (145-155). 139 
(110-165); BR= 1.9:1, 2.1:1; SP = 2,0: 1,5:1. 
1.6:0.9: 1. Tentacle sheaths sinuous to spirally 
coiled; TSW = 45.0-47.5, 45.0-50.0; TW basal 
45.0-50.0, metabasal 40.0-45.0, basal lentacular 
swelling absent. Prebulbar organs absent, 
muscular rings around basal pan of tentacle 
sheaths nol seen, retractor muscles originate at 
base of hulbs. Tentacular armature 
homeoacanthous. heteromorpbous, characteristic 
basal armature absent (Fig. 38). Hook form 
changes from compact, roscthorn-shaped 



66 



H. W. PALM & I. BEVERIDGE 




FIGURES 37 — 41 . Heteronybelinia pseudnmbusta sp. nov. from Lepidoirigla nwdesta. 37. Scolex of small form 
SAM AHC 28314. 38. Basal armature. 39. Metabasal armature. 40. Scolex of large form SAM AHC 28342. 41. 
Metabasal armature. Scale bars: Figure 37, 100 urn; Figures 38-39, 20 wn; Figure 40, 100 urn; Figure 41,15 \xm. 
FIGURES 42-44. Mixonybelinia cribhi sp. nov. from Hlaiycephaliis arcnarius Holotype, QM G 218047. 42. 
Scolex. 43. Metabasal armature. 44. Basal armature. Scale bars: Figure 42. 200 urn; Figures 43-44, 20 um. 



1 1 \ TACULARliD TRYPANORUYNCHS FROM AUSTRALIA 



Ii7 



(bothridial) to more slender hooks With stout base 
(anliboihridial) (Fig 39). Hook size in metabasal 
armature ranged between L =13.8-16.3. 
B = 11.3-13.8 (bothridial) and L= 163-17-5; 
B = 10.0-11.3 (antiboUiridial). hooks of basal part 
of tentacle smaller, between L - 10.0-11.0; 
B = 11.3-9.0 (bothridial) and L= 11.3-12.5, 
B = 8.8-10.0 (antibothridial). continuously 
increasing towards die tip; hsr = 6-7. 

iMria- fonn (2 paratypes): SL =1310 (Fig, 40); 
1 310, SW = 600, 620, pho = 810, 780, pv = 580, 
500; pb=390, 380; app = 330, 330; vel = 280. 
310. BL = 376 (360-390), 321 (310-330), 
BW = 124 (115-140). 121 (105-135); 
BR = 3.0:1; SP = 2.1:1.5:1. Tentacle sheaths 
sinuous to spirally coiled; TSW- 40-45. 42.5- 
47.5. TW basal 45.0-47.5. metabasal 30.0-35.0. 
30.0-35.0. Basal tentacular swelling absent, 
prebulbar organs absent, muscular rings around 
basal part of tentacle sheaths not seen. Retractor 
muscles originate at base ol bulbs. Tentacular 
armature homeoacanthous, heteromorphous, 
characteristic basal armature absent. Hooks 
change from compact and rosethorn-shaped 
(bothridial) to more slender hooks with a stout 
base (antibothridial) (Fig. 41). Hook size in 
metabasal armature between L= 12.5-14.5; 
B = ll.3-125 (bothridial) and L= 14.5-15.5; 
B =8.8-10.0 (antibothridial), hooks of basal pari 
of tentacle small, between L = 8.8-10:. B = 7.5- 
9.0 (bothridial) and L = 6.3-8.8; B = 6.3-7.5 
(antibothridial), increasing in size towards tip; 
hsr =6-7 

Remarks 

Heteronybelinia pseudorobusta sp. nov. is 
characterised by a heteromorphous tentacular 
armature consisting of small basal hooks 
mi teasing in size towards the metabasal region. 
The hook shape varies from rosethorn-shaped to 
slender hooks with a sioul base on different 
surfaces of the tentacle. Small basal hooks which 
gradually increase in size towards the metabasal 
armature are characteristic of Heteronybelinia 
mhusta (Linton, 1890). However, in contrast to 
// pseudorobusta sp, nov , //. robu.ua has minute 
tii\.il hooks, less than 5 in length, and the hook 
form is uniform along the tentacle. In die present 
specimens, the basal hook size is larger and the 
hook form is rosethorn-shaped. All other 
Heteronybelinia species differ in having a 
different hook shape or arrangement. Thus, the 
present specimens belong to a new species, H. 
pseudorubusta sp. nov., the name being derived 
from I he similarity in hook arrangement to H. 



robusta (Union, 1890). H, pseudorobusta is a 
species without a characteristic basal armature and 
with basal hook sites smaller than metabasal hook 
sizes, It therefore belongs to the Heteronybelinia 
species subgroup *Aa' in Palm (1999). 

Heteronybelinia pseudorobusta sp. nov. 
occurred in two different size groups, those 
smaller than I mm and those larger than 1.3 mm. 
respectively. Thus, die present description is an 
example of intraspeeific morphological variability 
within species of Heteronybelinia, apparently due 
to scolex contraction during fixation. 



Genus Mixonybelinia Palm. 1999 

Mixonybelinia bevtridgti (Palm. Walter. 
Scbwcrdtfcgcr & Rcimcr, 1997) 

Material exanunrd 

From Macruronus novaetetandiae (Hcctoi, 
1871): I postlarva. west coast of Tasmania, coll. 
K. Sewcll. June 1986 <QM C 218067) 

Ihuriptian 

SL = 3200. SW = 2750; pbo = 1750; pv = 850; 
pb - 1300; ppb = 80; vel = 950; app = 730; 
BL=1226 (1150-1310): BW = 290 <240-320|; 
BR = 4.2:1; SP = 1.3:0.7:1. Tentacle sheaths 
sinuous to spirally coiled; TSW = 100-130. 
Prebulbar organs absent, muscular rings around 
basal part of tentacle sheaths not seen. Retractor 
muscles originate in basal pans of bulbs Basal 
tentacular swelling absent; TW basal = 110-120. 
TW metabasal = 115-135. Metabasal armature 
homeoacanthous. heteromorphous, characteristic 
homcomorphous basal armature consisting of 
about 6-7 rows of slender hooks with elongated 
shaft, stout base and strongly recurved at op 
(L = 31 .3-40 0. B * 18.8-27.5). Metabasal hooks 
strongly recurved, rosethorn-shaped with large 
base on antibothridial surface: L = 50.0-52.5. 
B = 37.5— *2. 5, slender falcate hooks with stout 
base on bothridial surface I, = 55.0-60 0, 
B = 27.5-30.0; hsr = 6. 

Remarks 

The present specimen is characterised by a 
hoirieomorphous basal armature of 6-7 rows ol 
hooks and a heteromorphous metabasal armature 
of massive hooks, thus corresponding in form and 
size with specimens of M. bevehdgei described 
from African waters by Palm et al. 11997). Other 
features are the large scolex size as well as TSW 
and TW However, due to the contracted nature nl 



68 



H. W. PALM & I. BEVERIDCE 



the scolex of the current specimen, values for 
scolex and bulb proportions as well as the tentacle 
sheaths differed, indicating the variability of these 
characters due to the degree of contraction of the 
scolex. The present finding represents a new host 
and locality record for M. beveridgei. 



Mixonybelinia cribbi sp. nov. 
(Figs 42-^44) 

types 

Holotypc, posilarva from Platycephatus 
arenartus Ramsay & Ogilby. J 886, Heron Island, 
Queensland, coll. J. Reddick, Jan. 1997 (QM G 
21 8047). 

Material examined 
Types, 

Description 

Scolex craspedote, SL = 2920 (Fig. 42 i 
SW = 2200; pbo=l700; pv = 1020; pb=730; 
ppb = 220; vel = 760; app = 780; BL = 6 15 (600- 
630); BW = 232 (220-240). BR .= 2.7:1: 
SP = 2.3: 1.4:1. Tentacle sheaths short; 
TSW = 50-65. Prebulbar urgans absent, muscular 
rings around basal part of tentacle sheaths not 
seen. Retractor muscles originate in basal part of 
bulbs. Basal tentacular swelling absent; TW 
basal = 70-7.5, TW metabasal = 65-75 
Homcoacanihous, heteromorphous mciabasai 
armature consisting ol'roselhom-shaped hooks on 
botbridiaJ aud more slender hooks with stout base 
on antibothridial surface (Fig. 43); characteristic 
homeoinnrphous basal armature (Fig. 44) 
consisting of about 4 rows of slender hooks with 
enlarged base and strongly recurved at tip 
(L^ 12.5-17.5. B = 10 0-15.0 to 8.8-1 I 3) 
Metabasal hooks strongly recurved, rosethorn- 
shaped with large base on antibothridial surface- 
L = 20.0-22.5. B = 18.8-21.3; slender falcate with 
stout base along bothridial surface: L= 21.3-23.7. 
B = 15.0-17.5; hsr = 6-7. 

Remarks 

M. cribbi sp. nov. resembles M. beveridgei in 
scolex size as well as in tentacular armature Both 
species have a homeomorphous basal and a 
heteromorplious metabasal annature as well as a 
large scolex about 3 mm in length. However, A/. 
cribbi sp. nov. is clearly distinguishable from it. 
beveridgei by having only 4 rows of basal hooks 
compared with 6 in M, beveridgei. In addition, Ihe 
hooks have a distinctly enlarged base in M cribbi 



and all hooks are smaller (20-24 in M, cribbi 
compared with 46-68 in M. beveridgei, see Palm 
gl al, 1997). The new species was named alter Dr 
T. H. Crtbb, Department of Parasitology. 
University of Queensland, Australia. 



Mixonybelinia edninlinloni (Dollfus, 1960) 
(Figs 45-53) 

Material examined 

From Splnrna lewini (Griffith & Smith, 1843): 
10 adults from stomach. Flat Top island. 
Queensland, coll. B. G Robertson. 23.x. 1985 
(SAM AHC 28324); 1 adult, Geraldton. Western 
Australia, coli. B. G. Robertson, 27.xi.1986 (SAM 
AHC 28323). 

From Rhynchobaius djiddensts (Forsskal, 
1775): 9 postlarvae from spiral valve, Moreton 
Bay, Queensland, coll. S. Butler, 17 iv 1980 (QM 
G 218048-218060). 

From Carchartnnus melanopterus (Quoy <St 
Gatmard, 1824): 1 posilarva, (QMG4813). 

Description 

Adults: SL a 1780, 1620 (Fig. 45); SW= 1210, 
1 160: pbo = 1000, 1 000; pv = 760. 690; pb = 620, 
530: ppb = 60, 30; vel = 340, 330; BL = 585, 520, 
520. 530. 490, within single specimen 580-600; 
BW = 235 {230-240): BR a 2.5:1; SP = 1.6:1.2:1. 
Tentacles robust; tentacle sheaths straight. 
TSW = 50-90. Prebulbar organs absent, muscular 
rings around basal part of tentacle sheaths visible 
in some specimens Retractor muscles originate in 
basal part of bulbs: basal tentacular swelling 
absent; TW basal = 60-65; TW metabasal = 70- 
75. Mciabasai armature homcoacanthous, 
hcicromorphous (Fig. 46); characteristic 
homeomorphous basal armature consisting of 
about 10 rows of slender falciform hooks (Fig. 
46). Metabasal hooks strongly recurved along 
antibothridial surface" L = 20.0-25.0; B = 17.8- 
21.3; slender falcate hooks along bothridial 
surface. L = 32.5-35.0; B = 15.0-17.5. Basal 
hook size: L= 17.5-20.0, B = 10.0-13.8; hsr 
basal: 7-8: hsr metabasal: 8-9. 

Largest cesiode 53 mm, maximum width 880; 
560 segments; strobila craspedote, velum 
irregularly scalloped, up to 50 wide; pre- mature 
(no ovary) segments (Fig. 47) wider than long, 
100 x 1020 to 120 x 1020. Genital pores 
submargirml. in first third of segment; alternate 
irregularly. Cirrus sac elongate, 280-330 x 45-60 
(n=5) in mature segments, length:widtb ratio 
S 71 (4.7-7.1:1), distal pole directed 



THNTACULAR1ID TRYPANORHYNCHS FROM AUSTRALIA 



W 




46 





FIGURES 45-47. Mixonybelinia etlwinlintoni (Dollfus. I960,) from Sphyrna lewini. SAM AHC 28324. 45. Scolex. 
46. Basal and metahasal armature. 47. Premature proglottid. Scale bars: Figure 45. 150 urn; Figure 46, 25 pin; 
Figure 47, 60 urn. 



antcmmcdially, nearly reaching anterior end of 
segment; cirrus unarmed; internal and external 
seminal vesicles absent. Vas deferens coiled, 
extends to midline of segment, tlien posteriorly 
towards female, genital complex. Testes in two 
groups, not confluent posterior to ovary, 80 
(poral) and 116 (antiporal) (estimated), 30-40 in 



size (central), smallest testes at margin of medulla, 
externa] to osmoregulatory canal (15-25). 
distributed in 2-3 layers, extend anterior to cirrus 
sac; seminal receptacle present. Small vitelline 
follicles encircle medulla, between 24 longitudinal 
muscle bands. Osmoregulatory canals 5 and 20 in 
diameter. 



70 



H. W. PALM & I BEVERIDGE 












FIGURES 48-52. Mixonybelinut edwinUmoni (Dollfus. 1960) from Khynclwhatus djiddensis. QM G 218048-60 
48, Scolcx, 49, Basal armature, 50. Melabasal armature, 20 rows from the base. 51-52. Rulli with tentacle snealh. 
Scale bars Figure 48, 150 urn. Figures 49-50. 25 um. Figures 51-52, 50 urn. FIGURE 53. M. edwinlintoni from 
Carcharhinux melunoplenis. QM G 48 1 3. Scolcx Scale bar: 100 pirn FIGURE 54. Kotorella pmnn.mma (Stossich, 
1901) from Dasyatis fluvlotum. QM G 218063, Mature proglottid. Scale bar: 50 urn. 



TENTACULARIID TRYPANORHYNCUS FROM AUSTRALIA 



71 



Posdamae (from R. djiddensis): SL=16Q0, 
1500 (Fig. 48); SW= [000, 1180; pbo = 1020, 
950; pv = 680. 600; pb = 420. 470; ppb = 140, 
120; vel = 280, 250; app = 280, 250; BL«390 
(370-410), 430 (410-440); BW = 188 (180-190), 
ld3 (150-180); BR = 2.l:l, 2.6:1; SP= 2.4:1.6:1; 
2.0:1.3:1. Tentacle sheaths straight. TSW = 40- 
85. Prebulbar organs absent, museular rings and 
thickenings around basal part of tentacle sheaths 
visible in some specimens (Figs 5 1, 52). Retractor 
muscles originate in basal part of bulbs (Fig. 4K); 
basal tentacular swelling absent; TW basal = 80, 
80, TW metabasal = 70. 80. Homcomorphous 
basal armature of about 10 rows of slender 
falciform hooks (Fig. 49). metabasal armature 
homeoacanthous, heteromorphous (Fig. 50). 
Metabasal hooks strongly recurved along 
antibothridial surface: L = 22.5-25.0: B^ 17.5- 
20.0; slender falcate hooks along bothridial 
surface: L - 23 8-26 3; B = 12.5-15.0 Basal 
hook size: L = 17.5-20.0, B = 12.5-13,8; hsr 
basal: 7-8. hsr metabasal: 8-9. 

Postlan'a ( from C, melannpierus): SL = 1 648 
(Fig. 53); SW=848. pbo = 912, pv = 768, 
pb = 432, ppb = 96; app = 320; vel - 240; 
BL = 429 (411-443); BW = 171 (158-177); 
BR =2.5:1; SP = 2.1:1.8.1. Tentacles massive 
(TL= 630), TSW = 68-70, TW basal 77-80, TW 
metabasal 85-89, basal tentacular swelling absent. 
Prebulbar organs absent; muscular rings around 
basal part of tentacle sheaths not visible; retractor 
muscles originate at base of bulbs. Tentacular 
armature homeoacanthous, heteromorphous; 
characteristic basal armature absent. Hooks 
dimmish in size towards base; hook shape varies 
Irotri compact rose thorn- shaped (antibothridial) to 
slender rosethom-shaped hooks with stout base 
(bothridial) Hook size, metabasal, rows 16-17, 
L= 23.7-25.2; B = 14.2-15.8 (antibothridial) and 
L = 25-2-26.8; B = 9.5-12-0 (bothridial); basal 
L= 17.4-19.0; B = 14.2-15.8 (antibothridial) and 
L = 17.4-19.0; B= 12,6-14.2 (bothridial); 
hsr = 7-8. 

Remarks 

Palm and Walter (2000) redescribed 
Mixonyhelinia edwinlintuni (Dollfus. 1960) 
illustrating a distinctly different basal and 
metabasal armature within the species. While the 
metabasal armature has different hooks on the 
bothridial and antibothridial tentacle surface, the 
hooks of the basal armature appear to be 
homcomorphous. The arrangement of the hooks, 
together with their characteristic shape, and the 
scolex shape, indicate that the present specimens 



belong to M edwinlinioni. Rhynchahatus 
djiddensis and Corcharhinus melanopierus 
represent two new hosts for the species. 

Prior to the rc-dcscrip!ion of M. cdwinlintoni, 
Palm (1999) erected Heteronybetinia 
heleromurphi, a species which is also very similar 
to the present material. In H. heteromorpfii the 
basal armature is heteromorphous, while in A/. 
tdwtnltntoni it is homcomorphous. In addition, 
the type material of H. htterumvrphi was 
described as being acraspedote while M. 
cdwtfdinnmi is craspedote (Fig. 47). Interestingly, 
both species occur within the same host genus. 
Sphyma. Other characters such as the massive 
scolex form, the heteromorphous metabasal 
armature, and the testis sizes arc similar in both 
species. The type material of //. hererotrwrphi 
needs to be re-examined lo confirm Ihc 
differences between the species 



Mixonvbelinut southwelli (Palm & Walter, 
1999) 

Material examii\ed 

From Afakaira indica (Cuvicr, 1832): I 
posilarva. Cape Bowling Green, Queensland, coll. 
P. Speare, IK.i.v 1987 (QM G 2 1 806 1 >. 

From Chaerodon venustus (DcVts, 1885): I 
postlarva. Heron Island, Queensland, coll. R. 
Bray, 30L1991 (QM G 218062). 

Description 

SL = 2000, 1550; SW = 1050, 690; 
pbo = 1030, 890; pv - 850. 660; pb= 440, 590; 
ppb = 40; vel = 450. 220: app = 670, 270; 
BL = 41 1 (400-425), 548 (520-5S0); BW = 1 53 
(140-165), 153 (140-170); BR= 2.7:1. 3.6:1; 
SP = 2 3:19:1. 1,5:1.1:1 Tentacle sheaths 
spirally coiled; TSW = 55-60, 50-60 Prebulbar 
nrgans absent, muscular rings around the basal 
part of tentacle sheaths not seen: retractor 
muscles originate in basal pari of bulbs, 
tentacular swelling absent; TW basal = 50.0- 
55.0, 47.5-55.0, TW metabasal = 35^7, 30- 
35. Armature homeoacanthous. 

heteromorphous; characteristic basal armature 
consisting of about 14-16 rows. Antibothridial 
metabasal hooks massive, rosethorn-shaped, 
L = 15.0-17.5. 15.0-16.3. B = 13.7-15.0, 15.0- 
17.5; bothridial hooks more slender and slightly 
curved with stout base, L= 18 7-20.0, 17.5- 
20.0. B= II 3-12.5. 11.3-13.8. Basal armature 
homcomorphous, hooks falciform with stout 
base and strongly recurved tip. L= 15.0-22.5. 



72 



H. W. PALM & I. BEVERIDGE 



18.8-21.3; B = 7.5-8.7, 7.0-8.0; hsr basal = 6- 
7, hsr metabasal = 5-6. 

Remarks 

The postlarva from Chaerodon venustus, 
collected at Heron Island, is similar in scolex size 
and proportions, armature pattern and hook sizes 
to those described for M. southwelli by Palm and 
Walter (1999). Differences in scolex 
measurements of the specimen from Makaira 
indica are probably due to the shrunken condition 
of that specimen. The change in armature pattern 
from falciform basal hooks to heteromorphous 
rosethorn-shaped metabasal hooks is characteristic 
for M. southwelli. The distribution of the species 
is extended to eastern Australian waters, and 
Makaira indica and Chaerodon venustus represent 
new hosts for the species. 



Genus Kotorella Euzet & Radujkovic, 1989 

Kotorella pronosoma (Stossich, 1901) 
(Fig. 54) 

Material examined 

From Dasyatis fluviorum Ogilby, 1908: 2 
adults, Moreton Bay, Queensland, coll. S. Butler, 
26.ix.1980 (QM G 218063-218064). 

Description 

SL = 570, 660; SW = 200, 200; pbo = 400, 390; 
pv = 385, 510; pb = 80, 85; vel=110, 80; 
BL = 71 (70-73), 78 (71-88); BW = 39.4 (37.5- 
40.0), 40.0 (35.3-45.0); BR = 1.8:1, 2.0:1; 
SP = 5.0:4.8:1, 4.6:6.0:1. TW basal = 17.5, 22.5. 
Basal tentacular swelling absent. Tentacle sheaths 
straight; TSW = 1 1.3-15.0, 5.5-15.0. Prebulbar 
organs and muscular rings around basal part of 
tentacle sheaths absent; retractor muscles originate 
in basal part of bulbs. Tentacular armature 
homeoacanthous, heteromorphous; basal hooks on 
bothridial surface L = 6.3-7.5, 6.3-7.5, B = 5.0- 
6.0, 5.0-6.0, smallest hooks on antibothridial 
surface L = 2, increasing in size towards 
metabasal region. Small, incomplete worms to 
15 mm long, maximum width 570; 90 segments. 
Strobila acraspedote; pre-mature segments 30 x 
110 - 260 x 360, mature segments longer than 
wide or wider than long, 480 x 430 to 550 x 620 
(Fig. 54). In second specimen, mature segments 
500 x 440 to 550 x 650, and pre-gravid (with 
some eggs) segments 320 x 710 to 750 x 620. 
Genital pores submarginal, 210 from anterior 
margin; alternate irregularly. Cirrus sac elongate, 



150-250 x 30-60 (n = 5) in mature segments, 
length:width ratio 4.4:1 (4.0-5.0:1); cirrus 
unarmed; internal and external seminal vesicles 
absent. Vas deferens in large coils, extends to 
midline of segment, then posteriorly towards 
female genital complex. Testes 86 (81-100) in 
number, 45-85 in size (central), smallest testes 
20-50 (peripheral), distributed in single layer, 
confluent posterior and anterior to ovary. Ovary 
median, ovarian lobes 110-200 x 100-150; 
Mehlis' gland small, situated between ovarian 
lobes. No vitelline follicles in centre of the 
segment, follicles 33 (25—45) in diameter. No 
uterus observed, spherical eggs appear in 
parenchyma, egg diameter 22 (15-40). 

Remarks 

The morphology of Kotorella pronosoma 
(Stossich, 1901) is summarised by Euzet and 
Radujkovic (1989), Palm and Walter (1999) and 
Palm and Overstreet (2000). The present 
specimens from Moreton Bay correspond in 
scolex size and proportions as well as in the 
tentacular armature with the other described 
specimens. The sole difference noted was in testis 
number which exceeded 80 per segment in the 
specimens described. The species has previously 
been found in rays such as Aetobatus narinari 
(Euphrasen, 1790 ) (see Palm & Overstreet 2000), 
Himantura imbricata (Bloch & Schneider, 1801), 
Rhynchobatus djiddensis (see Shipley & Hornell 
1906; Palm & Walter 1999) and Dasyatis 
pastinaca (Linnaeus, 1758) (see Euzet & 
Radujkovic 1989; Palm & Walter 2000). Dasyatis 
fluviorum represents a new host record, and the 
range of distribution is extended to eastern 
Australian waters. The present finding suggests a 
cosmopolitan distribution for the species. 
Previously, the nearest known occurrence of this 
cestode to Australia was from the Java Sea under 
the name Nybelinia narinari (MacCallum, 1917), 
now a synonym of Kotorella pronosoma (see 
Palm & Overstreet 2000). 



Kotorelliella gen. no v. 

Diagnosis: Tentacles elongate, slender; retractor 
muscle originates at base of bulbs. Metabasal 
tentacular armature homeoacanthous, 
heteromorphous; hooks on bothridial surface 
uncinate becoming slender towards antibothridial 
surface. Basal hooks heteromorphous; 
characteristic basal armature present, arranged in 
a heteroacanthous atypica pattern with 



TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 



73 




FIGURES 55-61. Kntoreltiellajtmesi gen. ct sp. nov. from Taeniura lymma. Holotype, QM G 218065. 55. Scolex. 
56-57. Metabasal and apical armature, external view. 58. Basal armature, bothndial view. 59. BasaJ armature, 
antibothndial view. 6(M>1. Basal towards metabasal armature, external view. Note the interpolated hooks as in Fig. 
61. Scale bars: Figure 55, 150 urn; Figures 56-61, lOum. 



74 



H W. PALM & I. BEVER1DGE 



interspersed hooks on the antibothridial tentadt 
surface. Hooks solid. Slrohila unknown. 
Typc-spccics: Kn.jone:-i sp. no v. 



Kotorelliella jonesi sp, nov. 
(Figs 55-61) 

Typex 

Holotype, 1 posllarv3 from ihe spiral valve of 
Tuemura lymrna (Forsskiil, 1775), Heron Island, 
Queensland, coll. 1. Bevcndge & M. K. Jones. 
I l.vii.1998(QMG 218065). 

Material examined 
Holotype 

Description 

Scolex compact, 4 elongated bothridia, with 
hook-like microtriches along bothridial borders. 
Posterior margins of hothndia Tree, not fused with 
peduncle. SL = 1910 (Fig. 55); SW = 390; 
pbo = 800; pv = 1140; pb = 260; vel = 460; 
BL = 259 (2511-265). BW = 82 5 (80-8.5): 
BR = 3.1:1; SP = 3.1:4.4]. Tentacles long and 
slender. TW basal and metabasal = 21.2-23.8; 
TW apical = 17.5-20; basal tentacular swelling 
absent Tentacle sheaths straight; TSW = 15.0- 
17.7 Prebulbar organs and muscular rings around 
basal part of tentacle sheaths absent; retractor 
muscles originate in basal pari of bulbs. Metabasal 
(Fig. 56) and apical (Fig. 57) armature 
homeoacantlious, heteromorphous. Rosethom- 
shaped hooks with anterior extension of base 
cover bothridial tentacle surface (L= 13.5-15,0, 
B = 7.5-10.0); antibothridial surface with slender 
spiniform hooks, recurved at tip (L = 15.0-17.5, 
B = 5.0-6.3); hooks decrease in siic towards 
apical armature; hook shape remains constant 
(bothridial: L= 11.3-137, B= 100-11.2; 
antibothridial: L= 12.5-13.7, B = 4.5-5.5). Basal 
armature (Figs 58-61) with additional hooks 
interspersed on antibothridial surface; hook 
pattern heteroacanthous atypica, consisting of 
about 5 rows of hooks on bothridial (Fig. 59) and 
9-10 rows on antibothridial (Fig. 58) tentacle 
sutface; basal hooks on bothridial surface 
L = b. 3-10.0. B = 6.3-8.8. on antibothridial 
surface L = 2.0-10.0, B= 1.5-3.8. continuously 
increasing in size from base of tentacle towards 
metabasal armature (Figs 60-61) 

Remarks 

The holotype of Kotorelliella jonesi gen. et sp. 
nov. has a unique form of armature within the 



Tcntacuianidae, changing from a 
homeoacantlious. heteromorphous hook pattern 
with roseifturn shaped hooks in the metabasal 
armature to a heteroacanthous atypica hook 
pattern in the basal armature. The size of hooks in 
the basal and apical regions is smaller than in the 
metabasal region. 

On die basis of the scolex morphology. AY< 
jonesi sp. nnv appears to be similar to Kototella 
pronn:iima Both species have an elongated 
scolex, four elongated and slender hothndia with 
free margins, and short oval bulbs. In both 
species, the metabasal armature is 
heteromorphous, with hooks changing from 
uncinate on the bothridial surface to elongate on 
the antibothridial surface. Both species have a 
band of hook-like microtriches along bothridial 
borders, which are clearly visible using light 
microscopy Such a microthrix pattern is 
characteristic for tentaculariid trypanorbyneh 
cestodes. 

The basal armature of K. pronosoma is 
homeoacantlious, but that of Ko, jonesi sp. nov. is 
unusual, having additional hooks interspersed 
between the oblique spirals of hooks. Thus, Ihe 
basal tentacular armature is heteroacariihous 
atypica The basal region is homeoacanthous in 
Nybelinia, Heteronybelinia and Mixonybeliniu, 
bur in Teniacularia, the bothridial surfaces of the 
tentacle have extra rows of small hooks (see 
Beveridge & Campbell 1996). The unique feature 
of the tentacular armature suggests that the 
erection of a new genus is justified, even though 
it is known only from the larval stage. 

The new species was named after one of the 
collectors, Dr M. K. Jones, from the Centre for 
Microscopy and Microanalysis, University of 
Queensland, Australia. 



Unidentified material 

The following additional material, some of 
which represent new host records, was examined 
but could not be assigned to a species: 

Nybelinia sp. fTom the spiral valve of 
Rhynchobatus djiddensis. Fiat Top Island, 
Queensland, coll. B. G. Robertson, 25.x 1985 
(SAM AHC 28325) 

Nybelinia sp. from the spiral valve of 
Notorhynchus cepcdianus (Peron, 1807), south 
coast Kangaroo Island. South Australia, coll. B. 
G, Robertson. 10.ii.1985 (SAM AHC 28327) 

Nybelinia sp. from the stomach o! 
Notothynchus cepedianus. Young Rocks, 
Kangaroo Island. South Australia, coll. B. G. 



TENTACULARIID TRYPANORHYNCHS FROM AUSTRALIA 



75 



Robertson, 23 v. 1985 (SAM ARC 21354, 28328) 
Heteronybelinia sp, from the spiral valve of 
Carcharhinus amblyrhynchoides, Tommy Cut 
Mouth, Northern Territory, coll B. G, Robertson, 
19.ix. 1986 (SAM AHC 18326). 

Discussion 

The present study provides an overview for the 
first time of the tentaculariid trypanorhynch fauna 
Trom the Australian region. Three new species are 
added to the genus Nybelinia, two to 
Heteronybelinia and one to Mixonybelinia, with 
the latter two genera, as well as Kotorella, 
reported from the Australian region for the first 
time. In addition, a new genus KotarellicLia gen. 
nov., is erected, which is characterised by a 
homcoacanthous, heteromorphous mclabasal 
armature and a hclcroacanlhous basal armature. 
Other features such as the elongalcd, widely 



spaced bothridia and the short bulbs appear 
similar to Katorella. 

The tentaculariid trypanorhynch fauna in 
Australian waters is relatively species rich with 22 
(48 %) of the total of 46 known species occurring 
in these waters. Several of the new species may be 
endemic. However, the distribution of many 
species is still imperfectly known and a number of 
species previously thought to be endemic to a 
specific region have subsequently proven to be 
cosmopolitan (Palm et al. 1997; Palm 1999; Palm 
& Walter 1999, 2000). Reasons for broad 
distributions within tentaculariid trypanorhynchs 
may rcJate to life-cycle patterns, including widely 
distributed fish intermediate and elasmobranch 
definitive hosts. Another reason might be the low 
host specificity of the parasites, particularly in the 
intermediate host (Palm ct al. 1997; Palm & 
Walter 2000). 

II is difficult to clearly dislinguish between 



62 





FIGURES 62-63. Schematic drawing of scolex of Heternnybelinia pseuiltirnbutta sp. nov. from Lepidrilri gla 
mndptta, Illustrating (he arrangement of the dlfferenl tentacle surfaces wilhin lentaculuriid Irypanoihynehs. 62. 
View from bolhridial surface, with bothndial borders merging at the apex of the scolex. 63 View from external 
surface, with bolhridial borders widely spaced. Abbreviations, AB = anlibothndial view, B = bothndial view. 
Bo - bothridia, E = external view. I = internal view. 



76 



}[. W. PALM &. I BEVERIDOfc 



some tentaculanid species solely on Ihe basis of 
the tentacular armature (eg N. slrongyta), and 
additional taxonomic characters ate required. To 
date, the strohilae of only a Few tentaculanid 
species have been described in detail The 
present study demonstrates that, apart from the 
number and size or different genital structures, 
the cirrus sac length:width ratio might be a 
useful character as, for example, in H aunrah\\ 
sp. nov. and N. hemipristis sp. nov. The 
position of the anterior end of the ektus sac and 
the distance of the genital poie from the 
anterior end of the segment, as well as the 
number of layers of testis, whether the testes 
are confluent posterior to the ovary, and the 
number of testes anterior to the cirrus sac 
appear ro be useful taxonomic features. Further 
species descriptions are- needed to show whether 
or not these characters arc -variable, as is 
observed in a number of scolex characters, or 
whether (hey win be used more widely in 
tentaculanid systeroatics 

During this and previous studies, it hccauiL 
evident that the descriptions of the different 
tentacular surfaces of trypartorhyneh eestodes 
used by Dollfus (1942) and Campbell and 
Beveridge (1994) may be difficult to apply to 
larval lenlatulariids. In many cases it was hard 
to identify the various tentacle surfaces For a 
better understanding of the position of the four 
tentacles wiihin lentaculariid trypanorhynehs, 
the following two schematic illustrations are 
given (Figs 62. 63) as they may help to 
standardise the orientation of the different 
tentacle surfaces within tentaculariid 
trypanorhynehs The determination of tentacle 
orientation is based upon whether the 
bothridial borders merge at the apex of Ihe 
scolex (Fig- 62) or whether they are widely 
spaced (Fig. 63). The former figure shows the 
bothridial and the latter the external view of 
me tentacle surfaces. 

The description of Kown-UMlo jonesi gen. ct 
sp. nov. is of particular interest since its armature 
comprises both homeomorplious and 
heteromorphous components (Campbell & 
Beveridge 1994; Palm 1995. 1997). The species 
seems to be closely related to Kotorella 
pronosoma, which has a similar scolex 
morphology and metabasal armalure. Boih species 
have characteristic microtriches along the 
bothridial borders, which appear similar to those 
of Tentacularia caryplwenae and species of 
Nyliclinia (see Palm & Overslreel 2000) 
Although Beveridge et al. (1999) could not align 



Kotorella with the other tentaculariid genera using 
ciadistic analyses, we consider that Kotnrrlla 
pronosoma and Kotorelliella jonesi belong to Ihe 
Tenfaculariidne since they possess i 
humeoacanlhous metabasal armature and, more 
importantly, bands of hook-like microtriches 
along the bothridial borders. The latter character 
has as yet only been described for tentaculariid 
trypanorhynehs. 

Following the classification of Dollfus (l942i. 
only the metabasal armature is considered in 
classifying the tentacular armature. Howevet, in 
more leeent classifications, trypanorhynehs widi a 
distinct basal armature such as MLtodigma 
leptaleum Dailcy & Vogclbcin. 1974. 
Paroncomegas araya (Woodland, 1934) and 
Mixonybelinia beveridgei (Palm, Walter, 
Schwerdtfeger & Rcimer. 1997) have been placed 
in distinctive genera based in part on the basal 
armalure (Campbell & Beveridge 1994; Campbell 
etal. 1999; Palm 1999). Kotorclliella joncsi has 
thus been placed in a new genus. Interestingly, K. 
jom'si represents a transition from 
homeoacanthous towards heteroaeanihous 
armature as suggested by Campbell and Beveridge 
(1994, p. 57, lines 7-11). 

In summary, tentaculariid trypanorhynehs show 
considerable morphological variability, as can be 
seen by differences in scolex morphology between 
the genera Nybelinia, Heteronybelima and 
M irony helinia, and between Kotorella and 
Kotorelliella. The tentacular armature also differs 
between the species, varying from homeoacanthous 
and homeomorphous (Nybelinia. Afuonybelinia) to 
homeoacanthous and heteromorphous 
(Heteronybelinia. Kotorella, Mixonybelinia) 
pallerns. Kotorelliella has a homcoacantlious 
metabasal and a heteroaeanihous basal hook 
pattern. Additional methods, such as molecular 
gem-tic studies, might help to resolve not only 
species relationships but also the phylogeneti^ 
relationships of the species and species groups 
within this large trypanorhynch family. 



ACkNOWLLOOMUMli, 

Collecting was supported financially by the 
Australian Biological Rcsoutcc-s Study anil the 

Australian Restart* council Etnaiictal support was 

also provided hy the German Research Council | | 
PK b(4tl- 1 and 3 -2. We. wish to Hunk R A. Bray, S 
Butler, T M. Cribb , tit. K Jones, R. Martir, M 
O'CaJljghun, R. Norman, J. C. Pearson, S PicheUn. J 
Reddich, B. G Robertson , K. Sewell, P Speare and J 
Stevens for their contributions lo this studv 



TENTACULAR] LD TRYPANORHYNCHS FROM AUSTRALIA 



77 



ReniuiNCEs 



BEVERtDGE, I 1990. Taxonornic revision of 
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BEVER1DGE, I. & CAMPBELL, R. A 1996. New 
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BEVERIDCE, I., CAMPBELL. R. A. & PALM. H. W 

1999. Preliminary cladistic analysis of genera of (he 
order Trypanorhyncha Diesing, 1863. Systematic 
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CAMPBELL, R. A. & BEVER1DGE. I. 1994. Order 
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to the Cesiode Parasites of Vertebrates*. Eds L. F. 
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CAMPBELL. R.. MARQUES, F. & 1VANOV. V. 1999. 
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Eutelrarhynchidae) from the freshwater stingray 
Potamottygon mottiro in South America. Journal of 
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DOLLFUS, R.-P 1942. Etudes critiques sur les 
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DOLLFUS, R.-P, 1960. Sur une collection de 

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region de Dakar. Bulletin de I'lnsntut Francois 
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EUZET. L. & RADU3KOV1C, B. M. 1989. Kotorella 
pronosoma (Stossich. 1901) n. gen., n. comb., type 
des Kolorellidae, nouvelle famtlle de 
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Dusyatis posimaca (L., 1758). Annates de 
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FROESE. R. & PAULY. D. 1998. Fish Base y8 
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HEINZ, M. L. & DAILEY, M. D 1974. The 
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JONES. M. K. & BEVERIDCE, I. 1998 Nybelinia 
aueenslandcnsis sp n. (Cesloda: Trypanorhyncha) 
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Aiisnalia, with obscivarjon on the fine siruciurc of 
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KOROTAEVA. V D 1 97 1. fSome helminth fauna of 
the commercial marine fishes of the sub-order 
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Ii.vestiyo Tikhooneanskogo Nauchno- 



Issledovatel skogo hmititia Hybnogo Khozyaistvai 
Okeanografii 75: 69-84 In Russian. 

KOROTAEVA, V D. 1974. (Some helminth fauna of 
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Instituta ftybnoga Klwzyaistvai Okeanografii 88 
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LAST, P. R. & STEVENS. J. D. 1994. 'Sharks and 
Rays of Australia'. C.S.l.R.O : Australia. 

LESTER. R J. G., SEWELL, K. B., BARNES, A. &. 
EVANS, K. 1988. Stock discrimination of orange 
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PALM, H. W, 1995. Unicrsuchungen rur Systemalik 
von Russclhandwurmcrn (Cesloda: Trypanorhyncha) 
aus atlantischen Fischen. Berichie au.s dem Instilut 
fur Meereskunde, Kiel 275: 1-238. 

PALM, H. W. 1997 An alternative classification of 
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armature as being of limited importance. SystematU- 
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PALM. H. W. 1999. Nybelinia Poehe, 1926. 
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PALM, H. W. 2000. Trypanorhynch cestodes from 
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PALM, H. W. & OVERSTREET, R 2000. New records 
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Heteronybelinia palliata (Linton, 1924) comb, n. 
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PALM, H. W, He WALTER, T 1999. Nybetwa 
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PALM r H. W. &. WALTER, T 2000. Tentaculariid 
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PALM, H W., WALTER, T, SCHWERDTFEGER, G 
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Tetrarhynchen. Zoologische Jahrbucher 53: 559- 
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SEWELL, K. B. & LESTER, R. J. G. 1988. The number 
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VIJAYALAKSHMI, C, VIJAYALAKSHMI, J. & 
GANGADHARAM, T. 1996. Some trypanorhynch 
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13: 83-89. 



TWO NEW SPECIES OF THE GENUS LEPANUS BALTHASAR FROM 
SOUTH AUSTRALIA (COLEOPTERA : SCARABAEIDAE) 

E. G. Matthews & T. A. Weir 



Summary 

Ball-rolling dung beetles of the tribe Scarabaeini are recorded for the first time from South 
Australia. The new species Lepanus penelopae and L. loftyensis are described from southern Eyre 
Peninsula and the Mount Lofty Block respectively, and are compared with other species of the 
genus. There are brief remarks comparing Lepanus Balthasar with the related genera Sauvagesinella 
Paulian and Aptenocanthon Matthews. 



TWO NEW SPECIES OE THE GENES LEPANUS BALTHASAR FROM SOUTH AUSTRALIA 

(COLEOPTERA: SCARAB AEIDAE) 

E. G. MATTHEWS & T. A. WEIR 



MATTHEWS. E. G. & WBIR, T. A 2002 Two new species of Ihe genus Upturns Ballhasar 
from South Australia (Colcoptcra: Scaiahaeidae). Record* of the South Australian Museum. 
35(1): 79-84. 

Ball-rolling dung beetles of the tribe Scarabaeini are recorded for the first time from South 
Australia. The new species Lepanus penelopae and L. loftyensis are described from southern 
Eyre Peninsula and the Mount Lofty Block respectively, and are compared with other species 
of the genus. There are brief remarks comparing Lepanus Ballhasar with the related genera 
Sauvagesinella Paulian and Aptenocanthon Matthews. 

E. G. Matthews, South Australian Museum, North Terrace, Adelaide, South Australia 5000: 
amj T A Weir, CS1KO Division of Entomology, PO Box 1700, Canberra, Australian Capital 
Territory 260J. Manuscript received 8 August 200 J. 



At tlie time of the revision by Matthews (1974) 
of the Australian Scarabaeini, generally known as 
ball-rolling dung beetles, it was believed thai this 
group was absent from South Australia, although 
il was known from Victoria and Western 
Australia. However, in 1979 P. Greenslade 
obtained specimens of an undescribed species of 
the genus Lepanus Balthasar. which belongs to 
this tiibe. in the Marble Range of southern Eyre 
Peninsula; and in 1982 another undescribed 
species of the same genus was collected during 
extensive pitfall trapping undertaken by J and P. 
Greenslade in Kuilpo Kyeema Forest, southern 
Mount Lofty Ranges, in connection with a project 
to determine the effects of fire on the forest-floor 
fauna. Then in 1999 and 2000, more specimens of 
the second species were picked up by members of 
the Biological Survey of South Australia in the 
area of Mount Remarkable at the extreme northern 
end of the Mount Lofty Block environmental 
province, as well as near the Barossa Valley, using 
unbaked pitfall traps. Altogether, the two species 
are now known from five collection localities 
(Fig. 1), all situated on low mountain ranges at 
altitudes from 300 to just over 500 m. Kangaroo 
Island is part of the Mount Lofty Block but pitfall 
traps baited with human faeces, set throughout the 
island by one of the authors (EGM) in November 
1990, failed to collect any Lepanus. 

Lepanus is known from New Guinea, where it 
was first described, and from densely vegetated 
mesic and humid habitats along the northern, 
eastern and southern coasts of Australia as far 
west as the tingle forest near Walpole, Western 



Australia (Matthews 1974). The habitat is leaf 
litter, and food, where known, consists of 
vertebrate faecal matter. The descriptions below 
bring the total number of described Australian 
species of Lepanus to 23, but several undescribed 
species have been collected in recent years in 
eastern montane forests (R. Storey, pers. comm.). 

Specimens are deposited in the South 
Australian Museum, Adelaide (SAMA), the 
Australian National Insect Collection, Canbetra 
(AN1C), the Queensland Museum (QMBA); and 
the Queensland Department of Primary Industries 
collection at Mareeba (DPIM). 

All figures except the map arc by the senior 
author. 



Descriptions Of New Species 

Lepanus penelopae sp. nov. 
(Figs 1,2,3,5,7,9.12) 

Halotype 

'S. Aust. Eyre Pen. Marble Rge. Dense 
broombush. Pitfall. 4.10.79 PJ M. Greenslade', 
male, SAMA. 

Paratopes 

Same data as holotypc, I male and 5 females, 
AN IC, SAMA. 

Description 

Body uniformly piceous and nitid, legs rufous. 
Total length 5.0-6.0 mm. Maximum width across 



80 



E. G. MATTHEWS & T. A. WEIR 




FIGURE I . Known distribution in South Australia of species of Lepanus. ■ , L penelopae; • , L loftyensis. 



elytra 3.4-3.6 mm. Head - Dorsal surface even, 
very densely punctate with small shallow 
punctures not running together, some bearing 
short, very fine recumbent setae. Dorsal part of 
eye small, its maximum width contained about 25 
times in interocular distance. Prothorax -Anterior 
angles subquadrate. Sides of pronotum rounded, 
widest at broadly rounded posterior angles. Dorsal 
surface very densely and uniformly finely 
punctate, punctures separated by distances equal 
to a little more than their diameter, glabrous. 
Elytra - Striae shallowly impressed with regularly 
spaced moderately deep punctures separated by 2- 
4 diameters. Intervals nearly flat, smooth with 
very slightly uneven surface, glabrous, with 
minute punctures. Hind wings — Absent. Sterna - 
Meso- and metastema impunctate medially with 
large shallow cicatricose punctures laterally, 
glabrous. Legs - Protibia with three large teeth on 
outer edge, which is serrate between teeth and 
proximal to them. Claws (Fig. 7) a little expanded 
basally but not distinctly angulate or dentate. 
Abdomen - Pygidium (Fig. 9) with a very deep 




FIGURE 
I mm. 



Lepanus penelopae male, habitus. Scale line 



TWO NEW SPECIES OF LEPANUS BALTHASAR 



XI 






FIGURES 3-6. Tibiae of males in dorsal view. 3, L penelopae, left protibia; 4, L. loflyensh, right protibia; 5, L 
penelopae, right metatibin; 6. L iufiycnsis, light metalibia, b, brush of stiff setae; c, comb of flattened bristles. 
ScaJe line I mm. 



basal groove across whole width of disc, groove 
narrower and shallower in middle, surface inside 
it transversely striated Disc very finely punctate, 
glabrous, Aedeagus as in Fig. 12. Sexual 
dimorphism - Male with rounded expansion of 
inner apical end of protibia bearing apical comb 
of short wide flattened bristles (Fig. 3. c) and on 
lower surface a brush of long setae which is 
directed outwardly parallel to surface, extending 
beyond edge of comb (Fig. 3, b). Metatibia of 
male (Fig. 5) a little expanded at inner apical edge 
and bearing small apical tooth there. 

Remarks 

In the key to the species of Lepanus published 
by Matthews (1974), L. penelopae will go to 
couplet 16 because it has three protibial teeth, and 
then to couplet 18 because it has a transverse 
groove on the pygidium and glabrous dorsal 
surfaces. However, it does not have dentate claws 
as also required by this couplet. It can then be 
forced through couplets 19 and 20 to end up with 
L. illawarrensis Matthews of New South Wales, 
which appears to be the most closely related 
species, sharing many aspects of structure and 
secondary sexual characters. L. penelopae differs 



from L. illawarrensis in having simple claws, non- 
geminate elytral striae with much deeper strial 
punctures, the pygidial groove more strongly 
narrowed in the middle, the basal ridge of the 
pygidium straight in the middle, no trace of a 
median tubercle on the metasternum of the male, 
male hind tibia not twisted, and somewhat 
differently shaped aedeagal parameres. 

Etymology 

This species is named after Penelope 
Greenslade, who collected it as well as many of 
the other specimens of Lepanus described in this 
paper. 



Lepanus Inftvensis sp. no v. 
(Figs l,4,6,'8, 10, 11) 

Holutype 

'S. AUST. Pitfalls Kuitpo Kyeema Forest 15th 
March, 1982 P. Greenslade', male, SAMA. 

Paratopes 

Same data as holotype, 12 males, 4 females, 
ANIC, SAMA. 'S. Aust. Deep Ck Boat Hbr Lane 



82 



12. C. MATTHEWS & T. A WEIR 






9 



FIGURES 7-9. 7, I penelopae, claw segment of 
metatamis; 8, ditto, L loftyensh. Senile line 25 mnr. 
9. L penelopae, pygjdium nf male. Scale line 1 mm. 



turnoff pitfalls in heath 1-8 Dec 1983 P. 
Grecnsladc 1 , 1 female, SAMA. 'S AUST Ml 
Remarkable NP 11 km SSW Wilmington 
32M4'56"S 138"04-01"E 20-25 Nov 99 pitfall 
Flinders Ra. MAM00401', 5 females, ANIC, 
SAMA. \S. AUS. tl km SE Whispering Wall 
(near Para Wirra) 34"34'54"S 138 l) 55 , 12"E 
27-10-00 SMLR Survey BAR 03201*. 17 males, 
31 females, SAMA, QMBA, DPIM. 

Description 

Body uniformly fuscous to piccous, legs 
rufous. Total length 3.4-5.5 mm, maximum width 
across elytra 2.3-3.6 mm. Head - Surface even, 
very densely punctate with shallow coarse 
punctures running together in irregular transverse 
rows, bearing small recumbent setae. Dorsal pan 
of eye small, slit-like, its maximum width 
contained about 30 times in interocular distance. 



Prothorax - Anterior angles obi use. Sides of 
pronolum widest about one-quarter of the distance 
behind anterior angles, thereafter sliehily 
converging or subparallel to broadly rounded 
posterior angles. Dorsal surface very densely and 
uniformly punctate, punctures separated by 
distances equal to less than their diameter, bearing 
very fine, short, fully recumbent seiae, Elytra - 
Striae very superficial, geminate with slightly 
crenulate edges, impunelate Discal intervals Hat, 
in same plane as striae, with more or less undulate 
surface, finely shagreened, with a row of 
punctures bearing recumbent setae along edges of 
each interval, an additional row usually present in 
middle of intervals neat base. Hind wings - 
Atrophied, represented by short stubs, Sterna - 
Mesoslernum with shallow punctures on posterior 
half. Metasternum entirely densely punctate with 
shallow punctures laterally bearing short, fine 
recumbent setae. Legs - Protibia (Fig. 4) with 3 
large teeth on outer edge, which is serrate between 
teeth and proximal to them Claws (Fig. 8) 
strongly dentate. Abdomen - Pygidial disc simple, 
without groove, uniformly densely and sha] lowly 




%«^ w* 



FIGURE 10. Lepamis loftyensis male, habitus. Scale 
line 1 mm 



TWO NEW SPECIES OF LEPANUS BALTHASAR 



83 



punctate, punctures bearing very small recumbent 
setae usually worn off. Aedeagus as in Fig. 11. 
Sexual dimorphism - Male with rounded 
expansion of inner apical end of protibia bearing 
apical comb of short, wide, flattened bristles (Fig. 
4. c) and a brush of long setae underneath which 
is directed downward perpendicular to surface. 
Metatibia of male (Fig. 6) with strongly recurved 
apical end bearing acute inner tooth. Prothorax of 
male broader anteriorly, subquadrate; that of 
female slightly narrower anteriorly. Clypeal teeth 
of male slightly smaller than those of female. 

Remarks 

In the key to species of Lepanus in Matthews 
(1974) £. loftyensis will first go to couplet 16 
because it has three teeth on the outer edge of the 
protibia, then to couplet 17 because it has a simple 
pygidium and setose dorsal surfaces, then to L. 
villosus Matthews of north Queensland because 
of its crenulate elytral striae and densely punctate 
nieso- and mctasterna. It differs from L. villosus 
in having superficial elytra striae, reduced hind 
wings, very different parameres of the aedeagus, 
and being of much larger size. Further. L. 
loftyensis has Ihe setae on Ihe pronotum only 
about half the length of those on the elytra, the 
first elytral interval with a double row of seta- 
bearing punctures, and the upper edges of the 
epipleural carina not visible from directly above 
due to the curvature of the elytra. 

The designation MAM 00401 on labels of the 
Mounl Remarkable specimens refers to a locality 



which is described in Brandle (2001) as 
mountainous, on a hill crest, with loam/sand soil 
and an overstorey of Allocasuarina verticiltata at 
a cover density of 25-50%, and at an altitude 
above sea level of 430 m. For the Parra Wirra 
specimens the designation BAR 03201 refers to a 
ridge top covered with Eucalyptus fasciculosa I 
Acacia paradoxa woodland and an underslorey of 
Calytrix, Hakea and Xanthorrhnea, at an altitude 
of 515 m (L. Queale, pers. comm.). 

Etymology 

The name refers to the Mount Lofty Block 
environmental province in which all four 
collection localities of this species are contained. 
The block is Province number 3 as delineated tn 
the Biological Survey of South Australia, and 
extends from Kangaroo Island northward 10 
Mount Remarkable. The biota of this province is 
characterised as fully Bassian (Brandle 2001). 



Discussion 

The existence of the species here described as 
Lepanus loftyensis was briefly noted by Matthews 
(1984: 6, footnote) erroneously as an undescribed 
species of Sauvagesinella Paulian. Lepanus and 
Sauvagesinella are closely related genera which 
are not separable in ihe female sex, but in the 
male there are several secondary sexual characters 
which will distinguish them Unlike Lepanus, 
Sauvagesinella males have a prominent median 





11 



12 



FIGURES 1 1 & 12. Aedeagi in right and left views. 1 1. L loftyensis; 12. L penelopae. Scale line 1 mm. 



84 



E. G. MATTHEWS & T. A. WEIR 



lubercle on the metasternum (only a trace of a 
tubercle in some Lepanus), a row of close-set 
tubercles along the inner edge of the metatibia, 
and a more or less compressed metafemur with a 
longitudinal fold or ridge on its ventral surface. 
Sauvagesinella as understood here does not occur 
outside the extreme southern part of Western 
Australia, where there are three species (Matthews 
1974). 

Storey and Monteith (2000) mention that males 
of all species of Aptenocanthon Matthews, where 
known, have a similar form of the fore tibial apex 
with its inner angle expanded and bearing a short 
brush of stiff setae bent downwards at right angles 
to the upper tibial surface. This is a similar 
situation to that found in the two species of 
Lepanus described above (eg Fig. 3, b) and is also 
known to occur in several other species of 
Lepanus and all three species of Sauvagesinella, 
As well as this brush, there is a comb of flattened 
bristles at the protibial apex (Figs 3 and 4, c) 



which occurs, to varying degrees, in all known 
males of Aptenocanthon and Sauvagesinella as 
well as some species of Lepanus. Clearly, these 
male characters are not limited to Aptenocanthon 
but their value in classification must await further 
studies on the interrelationships of these genera of 
Scarabaeini, as also pointed out by Storey and 
Monteith (2000). 

In South Australia the only other known 
representatives of the dung beetle subfamily 
Scarabaeinae belong to the tribes Onthophagini 
(native and introduced Onthophugus Latreille), 
Onitini (introduced Onitis Fabricius) and 
Oniticellini (introduced Euoniticellus Janssens), 
none of which make or roll food balls. All these 
groups are differentiated from the tribe 
Scarabaeini by their middle and hind tibiae, 
which are strongly expanded apically. In 
Scarabaeini these tibiae are slender and only a 
little expanded, an adaptation for rolling the food 
material. 



References 



BRANDLE, K. 2001. A biological survey of the 
Flinders Ranges, South Australia 1997-1999. 
Biodiversity Survey and Monitoring, National Parks 
and Wildlife, South Australia, Department for 
Environment and Heritage, pp. i-xviii, 1—455. 

MATTHEWS, E. G. 1974. A revision of the scarabaeine 
dung beetles of Australia. II. Tribe Scarabaeini. 
Australian Journal of Zoology, Supplementary Series 
24: 1-211. 



MATTHEWS, E. G. 1984. A guide to the genera of 
beetles of South Australia Part 3. South Australian 
Museum. Special Educational Bulletin Series 6: 1- 
60. 

STOREY, R. 1. & MONTEITH, G. B. 2000. Five new 
species of Aptenocanthon Matthews (Coleoptcra: 
Scarabaeidae: Scarabaeinae) from tropical Australia, 
with notes on distribution. Memoirs of the 
Queensland Museum 46(1): 349-358, 



A CHECKLIST OF CISSEIS (SENSU STRICTO) GORY & LAPORTE, 1839 
(COLEOPTERA : BUPRESTIDAE : AGRILINAE) 

Shelley Barker 



Summary 

A checklist of Cisseis (s.s.) Gory & Laporte, 1839 is presented and discussed, summarising the 
information available on the genus at the present time. 



A CHECKLIST OF CISSEIS (SENSU STRICTO) GORY & LAPORTE, 1839 
(COLEOPTERA: BUPRESTIDAE: AGRILINAE). 

SHELLEY BARKER 



BARKER, S. 2002. A checklist of Cisseis (sensu siricto) Gory & Laporte. 1839 (Coleoptera: 
Buprcstidae: Agrilinae). Records of the Souih Australian Museum 35(1): 85-90. 

A checklist of Cisseis (.«.,«.') Gory & Laporte, 1839 is presented and discussed, summarising 
the information available on the genus at the present time. 

S. Barker, Department of Entomology, South Australian Museum, North Terrace. Adelaide, 
South Australia 5000. Manuscript received 3 December 2001. 



Material 

Specimens examined came from the following 
institutions: 
AMSA - Australian Museum, Sydney 

AN1C - Australian National Insect Collection, 
CS1RO, Canberra 

BMNH - The Natural History Museum, London 

BPBM - B. P. Bishop Museum, Honolulu 

HMOE - Hope Museum, Oxford 

HUMB - Humboldt University Museum, Berlin 

MCSNG - Museo Civico di Storia Natiirale 
Genoa, Italy 

MNHN - Museum National d' Histoire Naturelle, 

Paris 

NMVA - National Museum of Victoria, 

Melbourne 

RMCE - National Museum of Czech Republic, 

Prague 

QMBA - Queensland Museum, Brisbane 

RMBB - Royal Museum. Brussels 

SAMA - South Australian Museum, Adelaide. 

Abbreviations: 
WA - Western Australia 
NWA - Kimberly area. Western Australia 
Q - Queensland 
NSW - New South Wales 
V - Victoria 
SA — Soulh Australia 
T - Tasmania 
NT - Northern Terriloi v, 



iNITtODUCTlON 

Many of the Australian bupresttd genera have 
not been reviewed since the death of H. J. Carter in 
1941. Because of the large amount of subsequent 
collecting and confusion with species 
identification, the genera are in a parlous state of 
order. One of worst has been Cisseis (Coleoptera: 
Buprestidac: Agrilinae), last researched by Carter 
(1923, 1929). With the intention of revising 
Cisseis, the author commenced work on the group 
in 1997. All of the readily available types, more 
than half of which are housed in European 
museums, have now been examined. Some of the 
difficulties have been resolved and a number of 
new species described (Barker 1998, 1999a, 1999b, 
2001). Some species clearly did not belong in 
Cisseis and these, together with nine new species, 
were placed in a new genus Stanwatkinsius Barker 
& Bellamy (2001), Carter (1923) did not examine 
male genitalia, which the author has found to be 
highly diagnostic. As a result, many of his 
specimens located in various museum collections 
are incorrectly identified. In order to summarise the 
knowledge available on the genus, a checklist has 
been constructed of valid species together with 
their distributions by state or country. Types and 
their locations are also listed The following 
abbreviations are used: HT = holotype; ST = 
syntype; LT = lectotype; AT = allotype. 

Checklist of Cisseis (sensu stricto) Gory & 
Laporte, 1839. 

uberrans Barker, 2001. Trans. Rov. Sac. S. Anst. 
125: 98. (cT HT, Woodridge, WA, H. 
Demarz, SAMA I 21504). WA. 



86 



S. BARKER 



acuducta (Kirby, 1836)(7>m-/7v.?) Faun, Bor. 

Amen; 162 (d HT. N Scotia, BMNH). 

Q, NSW.V.SA.T. 
marmorea (Gory & Laporte, 1839)(£r/itiK). 

Mon, Bupr, ii: 4. (HT, MNHN not seen) 
cuprifrons Kerremans, 1898. Ann Soc. Em. 

Belg. 42: 157. (d HT. Australia. 

Standing. BMNH) 
lacta Kerremans, 1903, in WytSmun Genera 

Ins. fas. 12: 227. (d HT, Australia, 

Moffarts. BMNH) 
adusla Barker. 2001. Trans. Roy. Soc. S. Aust. 

125: 101. (d HT. Lucindale, SA, 

Feueheerdt, SAM A ! 21505). SA. 
albertisi Gestro, 1877. Ann. Mus. Civ. Sior. Nat. 

Genova. 9: 357. (3 ST sex unknown, 

Somerset, Australia, D'Albertis, 1/75 

MCSNG: 9 ST. Somerset, Australia, 

RMBB). Q. 
albasparsa Gory & Laporte, 1839. Monograph ii: 

3. (HT, MNHN not seen). NT, Q. 
cuprivenlris Kerremans, 1898 Ann. Soc. Ent 

Belg. 42; 161. (9 HT, Australia, 

Chevrolat. BMNH). 
inflammata Carter, 1923. Proc. Linn. Snc. 

N.S.W. 48: 167. (d HT. S. Johnstone R., 

Q, H. W. Brown AMSA K67240) (new 

syn.). 
u</uilonia Bellamy. 1991. Tijdschr v. Ent. 134: 

171. ((J HT, BPBM not seen). Philippine 

Arch. 
urmstrnngi Barker, 2001. Trans. Roy. Soc. S. 

Am. 125: 101. (d HT, BoganR, NSW, 

J. Armstrong, NMVA). NSW. 
augustgoerlingt Barker, 2001. Trans. Roy. Soc. S. 

Aust. 125: 102. (d HT, Marloo Stn., 

Wurarga, WA, A. Goerling. AN1C). WA. 
autocyanea Carter. 1934. Proc. Linn. Soc. N.S.W. 

59: 258. (9 HT, Fletcher, Q, E. Sutton. 

AMSA K67296). NSW. Q. 
bedfordi Obcnberger, 1923. Casopis Cs Spot 

Enlom. 32: 9. (9 HT, Cape Bedford, Q, 

NMPC). Q. 
brooksi Barker. 2001. Trans. Roy. Soc. S. Aust. 

125' 102 (d HT, Marecba, Q, J. G. 

Brooks, ANIC). Q 
broumensis Barker, 2001. Trans. Roy. Soc S 

Aust 125: 103. (d HT, Broome, WA, H. 

W. Brown, SAMA 1 21509). NWA. 
browni Carter, 1934. Proc. Linn. Soc. N.S.W. 59. 

258. (d HT, Dcdari, WA. H. W. Brown. 

AMSA K67297). WA. 
airttrcBa Obenberger, 1935. Col. Cataiogus 12: 

846. (9 IT. Herherton, 0. C. J. Wild, 

QMBAKrcplac. name). Q. 



elliptica var frontalis Carter, 1923. Proc. 

Linn. Soc. N.S.W. 48: 170. (homonym). 
carteri Obenberger. 1924, Arcluv, (lir Namrg. 90: 

109. \6 LT. Yilgarn, WA, NMPp.WA. 
chalcophora Barker, 2001. Trans. Roy. Soc. S. 

Aust 125: 103. («J HT, Kapunda, SA, S. 

Barker, SAMA I 21507). SA, 
cfermonti I'hcrj', 1945. Bull. Soc. Ent. Fr. 50: 46 

(HT, MNHN not seen). Bougainville. 
corpulenta Barker, 2001 Trans. Roy Soc, S. 

Aust, 125: 104. (<J HT, Tallering sin., 

Pindar, WA, S Barker, SAMA 121511). 

WA. 
cupreicollis (Hope, \Mf>)(Ethon). Ann. Mag, Nat. 

Hist. 17; 64. (6 HT, Moriatta (sic), SA, 

HMOE no. 893) SA 
aenicollis (Hope, \846)(Etfion), Ann Mag. 

Nat. Hist. 17: 65. (9 HT, Adelaide, 

HMOE no. 892 2/2). 
cupreala Barker, 2001. Trans. Ray. Soc. S. Aust. 

125: 105. (d HT, Ml Spec, Q, J. G. B., 

ANIC). SA, NSW. Q. 
cupripennis (Cnerin, \%>2>b)(Buprestis). Voy. 

Coauille- 65. (HT, MNHN not seen). 

NSW, V. 
semiscabrosa Thomson, 1879 Typi Bupr. 

Mus. Thomson. App 1A' 53. (d HT, 

Sydney Knew syn.). 
niiidicotlis Kenetnans, 1898. Ann. Soc. Ent. 

Belg. 42: 162. (d HT, Auslralie, Bales, 

BMNHHnew syn.). 
cyanea Barker, 2001. Trans. Ro\, Soc. S. Aust. 

125: 105, (d HT, Wialki'WA, S. Barker, 

SAMA I 21512). WA. 
cyaneopyga Carter, 1923. Proc Linn Sac. N.S.W 

48- 170. (2 LT, Lake Austin, WA, H. 

W. Brown, AMSA K67292) WA. 
gocrlingi Carter, 1936 Proc. Linn. Soc. 

N.S.W. 61: t04. (2 HT, Marloo sin, 

Wurarga, WA, A. Goerling, AMSA 

K67466)(new syn.). 
cyanura Kerremans, 1898. Ann. Soc, Ent. Belg. 

42: 163. (2 d 6 ST & 2 ST BMNH). Q. 
derbyensb Barker, 2001. Trans. Roy. Soc S. Aust 

125: 106. (d HT. Derby, H. W. Brown, 

SAMA I 21 520). NWA. 
duodecimmai-nlata (Fabricius, \XQ\){Buprestis). 

Sys. Eleuth.: 191. (HT sex unknown, 

BMNH). All mainland states. 
duodecimguttata (Boisduval, 1835) 

(Buprestis). Vov. Astrolabe, Enlom. 2: 

93. (HT, MNHN not seen). 
quaturodecimnaiata Hope, 1846. Ann. Mag 

Nat. Hist. 17: 64. (HT not seen. HMOE). 
pusiitlata Thomson, 1879. Typi Bupr. Mus. 



CHECKLIST OF CISSHtS (S.S. ) GORY & LAPORTE, I &39 



87 



Thorns. App 1A: .51 (HT sex unknown, 

MNHN). 
biologia Froggatt, 1910. Australian Insects: 

165. (HT not seen, AMSA?)(ncw syn.). 
fallaciosula Obcnbergcr, 1935. Acta. Soc. 

Ent Csl 32: 36. (HT sex unknown, 

PMCE )(ncw syn.). 
etoMgatula Blackburn, 1888. Proc. Linn. Soc. 

N.S. W. 3; 891 ( tS HT, NT. SAMA). NT. 
elliptka Carter, 1923. Proc: Linn. Soc. N.S.W. 48: 

170. (9 L.T, Cue. WA, H. W. Brown. 

NMVA). WA. 
excelsior Barker, 2001. Trans. Ru\. Soc S. Ausr. 

125: 107. (d HT, Mt Carbine, 0, T. M. S. 

Hanlon & M. Powell, SAMA 1 21514). Q. 
ernestadamsi Barker, 1999. Rec. S. Aust. Mas. 32: 

46. (d HT. Edungalba. Q, E. E. Adams. 

ANIC). Q. 
Jascigera Obenbergei, 1919, Ent, Miiteilungen 8: 

20. (d HT, New Guinea, PMCE no 

23772). New Guinea 
fraterna Kerremans, 1900. Mem Soc- Enr. Belg. 

7- SO (2 HT, Germ N. G., Webber. 

BMNH). New Guinea. 
frontalis Kerremans, 1898. Ann. Soc: Ent. Belg. 

42: 157. (d HT, New Gumee, Standing. 

BMNH). New Guinea. 
fulgidicollis Maclcay. 1888. Proc. Linn. Soc. 

NSW. 3 1231 (2 6<S ST, Derby, 

NWA, ANIC). NWA, Q. 
gibbera (Carter. \9TI)(Neospades). Irons. Rov. 

Soc. 5 Aust. 61: 121. (2 HT, Mullaly, 

NSW, H. J. Carter, AMSA Kl 11950). 

NSW 
gouldi (Hope, 1&46)( Erhon). Ann. Mag. Nat. Hist. 

17: 65. (9 HT. Port Essington. HMOE 

no. 889). NT 
Iteroni Carter, 1934. Proc. Linn. Soc. N.S.W 59 

259. (d HT, Dorrigo. NSW, W. Hcrron, 

AMSA K67298). NSW. 
impressicollis Macleay, 1878. Enr. Soc. N.S.W. 2: 

248. (9 HT. Gayndah, AMSA K327I3). 

Q. 

viridiaurea Maclcay, 1878. Ent. Soc. N.S.W. 

2; 248. (2 2 2 ST. Gayndah, AMSA no. 

K327l2)(ncwsyn.). 
viridkuprea Kerremans, 1 898. Ann. Sot: Ent. 

Belg. 42: 160. (9 HT, Cocktown (/lie), 

Standing, BMNHHnew syn.). 
nitida Kerremans, 1S9S. Ann. Soc Ent. Belg. 

42: 164. (d HT, Cocktown (sir), 

BMNH)(new syn.). 
inops Kerremans. 1898. Ann. Sue. Ent. Belg. 42: 

168. (9 HT, Australie. Standing, 

BMNH). NSW. 



kohouli Barker, 2001. Trans. Rov. Soc. S Aust. 

125: 107. (d HT. Canberra, K. Pulien, 

ANIC). NSW, Q, 
lalicollis Carter 1923. Proc. Unn. Soc N.S.W. 

48: 169 (2 HT, BMNH). Q, 
leucoslicta (Kirby, \%W)(Buprestis\. Trans. Enr. 

Soc. Lond. 12: 382. (9 HT, BMNH). All 

mainland states. 
attrulenia Kerremans, 1898. Ann. Soc. Ent. 

Belg. 42: 159. (9 HT, Australia, 

Chevrolat, BMNHHnew syn.), 
juigidifrons Kerremans. 1898. Ann. Soc. Ent. 

Belg. 42: 161. (2 HT. Australia, 

Chevrolat. BMNH). 
macmillani Barker, 2001. Trans. Rox. Soc. S. 

Ausr. 125: 108. id HT. Wanneroo. WA, 

R. P. McMillan. SAMA 1 21516). WA. 
macqueeni Barker, 2001 Trans. Roy. Soc S. 

Aust 125: 108. (6* HT, Milmcrran. O. J. 

Macqueen, ANIC) Q 
maculata (Gory & Laporte, )B39)(Ethon). Man 

Bupr n 5, (HT, MNHN not seen). NSW 
marmnrata Gory & Laporte, 1839. Man Bupr. ii: 

4. (HT.MNHN not seen). NSW, Q. 
viridicollis Thomson, 1879. Tvpi Bupr. Mus. 

Thomson. App. 1A. 50. (d HT, MNHN). 
aena Kerremans, 1898. Ann. Soc. Em. Belg. 

42; 158. (3 2 2 ST, Sydney, Bates. 

BMNH). 
minutissima Thomson, 1879. Typi Bupr. Mt»s, 

Thomson. App. 1A. 54. (HT, MNHN not 

seen). SA, V 
simplex Kerremans, 1898. Ann. Sac. Ent. 

Belg. 42: 171. (2 d d ST & 2 ST, 

Australie, Standing, B.MNH)(nc-w syn.). 
modesta Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 

170. (d HT, Australia, Stark, Chevrolat 

collection, BMNH). WA. SA. V. 
myallae Carter, 1934. Proc Linn. Soc. N.S.W. 59: 

260. (2 HT, Bogan R., NSW, ) 

Armstrong. AMSA K67299). NSW. 
nigrita Kerremans, 1898.A«/i Soc. Ent. Belg. 42; 

168 (d HT, Australie, Fairmairc. 

BMNH) 
nigripennis Macleay, 1888. Proc Linn. Sm 

N.S.W. 3: 1231. (d HT, Barnor (sir.) 

Range, NWA. Froggeli, ANIC). NT, WA. 
ignicolhs Kerremans. ISt'S. Attn. Soc Ent. 

Belg. 42: 164. (6 ST, Australie. 

Standing, BMNH; S ST, Australie. 

Chevrolat, BMNH; 2 o" 6 ST, Port 

Dcnison BMNH Knew syn.), 
tiigromaculata Kerremans, 1895. Ann. Soc Ent. 

Belg. 39: 218, (HT not in BMNH). 

Malacca. 



S. BARKER 



nitidiventris Carter, 1934. Proc. Linn. Soc. 
N.S.W. 59: 260. (2 HT, Gosford, NSW, 
N. MacGregor, AMSA K67300). NSW, 

Q- 
niveosparsa (Carter, \927)(Neospades). Proc. 

Linn. Soc. N.S.W. 52: 228. (8 HT, Bogan 

R., NSW, J. Armstrong, AMSA 

K67288). NSW, SA. 
notulata (Germar, \M8)(Ethon). Linn. Ent.: 178. 

(8 HT, Adelaide, HUMB). SA, NSW, T. 
atroviolacea Thomson, 1879. Typi. Bupr. 

Mus. Thorns. App. 1A: 52. (8 ST & 2 

ST, Austr, MNHN)(new syn.). 
morosa Kerremans, 1898. Ann. Soc. Ent. 

Belg. 42: 158. (2 8 8 ST, Australie, 

Stark, BMNH)(new syn.). 
nubeculosa (Germar, 1848)(£7/zon). Linn. Ent.: 

176. (5 LT, Adelaide, HUMB). SA, V. 
chalcoptera (Germar, 1848)(£//ion). Linn. 

Ent: 177. (2 8 8 ST, Adelaide, HUMB). 
similis Saunders, 1871. Cat. Bupr.: 103. (2 

HT, Adelaide, BMNH)(new syn.). 
oblonga Kerremans, 1903. in Wytsman Genera 

Ins. fas. 12: 229. (2 HT, Australie, 

Fairm., BMNH). V, NSW, Q, T. 
obscura Blackburn, 1887. Trans. R. Soc. S. Aust. 

10: 252. (2 HT, T 320, type locality 

unknown, BMNH)(unique). 
opima Thomson, 1879. Typi Bupr. Mus. 

Thomson. App. 1A: 50. (2 HT, 

Champion Bay, MNHN). WA. 
oralis Carter, 1923. Proc, Linn. Soc. N.S.W. 48: 

170. (2 HT, Nov. Holl. occid.. Fry Coll. 

1905.100, BMNH 37811)(unique). WA. 
parva Blackburn, 1887. Trans. R. Soc. S. Aust. 

10: 253. (<J HT, BMNH). WA, SA, V, 

NSW. 
pygmaea Blackburn, 1891. Trans. R. Soc. S. 

Aust. 14: 299. (2 HT, Victoria, C. 

French, NMVA)(new syn.). 
patricia Carter, 1935. Proc. Linn. Soc. N.S.W. 55: 

180. (8 HT, Bunbury, WA, F. L. 

Whitlock, AMSA K67302). WA, V, 

NSW. 
pauperula Kerremans, 1898. Ann. Soc. Ent. Belg. 

42: 167. (<J HT, Australie, A. Deyr., 

Chevrolat collection, BMNH). T. 
prasina Carter, 1923. Proc. Linn. Soc. N.S.W. 48: 

168. (8 LT, Australia, Blackburn's 

collection, 3267, SAMA). SA, Q. 
puella Kerremans, 1898. Ann. Soc. Ent. Belg. 42: 

170. (2 8 8 ST & 2 ST, Australie, 

Fairm., BMNH). Q. 
curta Kerremans, 1903. in Wytsman Genera 

Ins. fas. 12: 229. (8 HT, Peak Downs, 



Fairmaire, Mus. Godeffroy no. 11142, 

BMNH). 
pulchella Carter, 1923. Proc. Linn Soc. N.S.W. 

48: 171. (HT sex unknown, French 

collection, NMVA). Q. 
pulleni Barker, 2001. Trans. Roy. Soc. S. Aust. 

125: 109. (8 HT, Edungalba, Q, S. 

Barker, SAMA I 21518). ACT, Q. 
regalis Thomson, 1879. Typi Bupr. Mus. 

Thomson. App. 1A: 50. (8 HT, 

Queensland, MNHN). NT, Q. 
robertfisheri Barker, 1999. Rec. S. Aust. Mus. 32: 

47. (8 HT, Melrose, SA, R. H. Fisher, 

SAMA I 21407). SA. 
roseocuprea (Hope, \846)(Ethon). Ann. Mag. 

Nat. Hist. 17: 64. (2 HT, Moriatta (sic), 

SA, HMOE no. 889). WA, SA, V, NSW. 
dispar Blackburn, 1891. Trans. R. Soc. S. 

Aust. 14: 297. (2 HT, 3832, BMNH). 
fairmairei Kerremans, 1898. Ann. Soc. Ent. 

Belg. 42: 169. (2 HT, Australie, 

Fairmaire, BMNH). 
cuprca Kerremans, 1903. in Wytsman Genera 

Ins. fas. 12: 228. (2 HT, NSW, Standing, 

BMNH). 
rubicunda Kerremans, 1898. Ann. Soc. Ent. Belg. 

42: 169. (8 HT, Australie, Fairm., 

BMNH). WA, SA, V, NSW, Q. 
undulata Kerremans, 1903. in Wytsman 

Genera Ins. fas. 12: 227. (2 HT, NSW, 

Standing, BMNH)(new syn.). 
purpurea Kerremans, 1903. in Wytsman 

Genera Ins. fas. 12: 228. (2 HT, Sidney 

(sic), Fairm, Mus. Godeffroy no. 

11108)(newsyn.). 
scabiosa (Boisduval, ]835)(Buprestis). Voy. 

Astrolobe Entom. 2: 96. (HT, MNHN not 

seen). 
scabrosula Kerremans, 1898. Ann. Soc. Ent. Belg. 

42: 167. (3 8 8 ST & 2 ST, Australie, 

Fairm., BMNH). NSW, Q. 
semiobscura Kerremans, 1898. Ann. Soc. Ent. 

Belg. 42: 170. (8 ST, Australie, Fairm., 

BMNH; 2 22 ST, Australie, Deyr., 

BMNH). SA, V. 
septuosa Barker, 2001. Trans. Roy. Soc. S. Aust. 

125: 109. (8 HT, Charity Ck Bridge, 

Manning R., NSW, S. Watkins. SAMA I 

21521). SA, NSW, Q. 
sexnotata Fauvel, 1891. Rev. d'Ent. 10: 181. (HT, 

MNHN not seen). New Caledonia. 
signaticollis (Hope, l&46)(Ethon). Ann. Mag. Nat. 

Hist. 17: 64. (8 HT, Port Essington, 

HMOE no. 891 ). WA, NT, Q. 
speciosa Barker, 2001. Trans. Roy. Soc. S. Aust. 



CHECKLIST OF CISSI-JS (S.S J GORY & LAPORTE, 1839 



J25: 110. U-i HT, Midland Jem. WA, R 

P McMillan, SAMA I 21523) WA 
stellata Barker, 2001 Ttms Ray. Soc. S. Aust. 

!25: III. (6 HT, Cairns. SAMA T 

21524). 0, 
stigmata Gory & Laporte, 1 839. Man. Bupr. ii: 3. 

(HT. MNHN not seen). WA. 
subbifascialis Carter, 1927. Proc. Linn. Soc. 

N.S.W. 52: 229. (9 HT. Bogan R_. NSW. 

J.A..AMSAK0729I). NSW. 
tastnatuea Kerremans, 1898. Ann. Soc. Ent. Rely,. 

42: lfi5. (2 ST, Tasnunic, Obcrthur, 

BMNH: ST. Tasmanie. Chcvrolat. 

BMNH).T, V, NSW. 
trimentula Barker, 2001. Tram. Roy. Soc. S, Au.it. 

125: 111. (d HT, Pilliga East SF, NSW. 

S. Watkins, SAMA I 21 526). V. NSW. Q. 
tyrrhena Carter, 1923 Proc. Linn. Soc. N.S.W 

48: 168. (9 HT. Kalamunda. WA. H M. 

Giles, AMSA K67293). WA 
\icina Kerremans, 1898 Ann, Soc Ent Belg 42: 

164. (<J HT. Australie, Standing, 

BMNH). V, NSW, Q. 
collaris Kerremans, 1903. in Wytsman, 

Genera his. fas. 12: 229. (2 d 6 ST, 

Cayndah, Fainnaire, BMNH). 
or naia Kerremans, 1903- in Wytsman Genera 

Ins. fas. 12: 229, (6 HT, NSW. Standing, 

BMNH). 
v'uiUtcea Kerremans, 1903. in Wytsman Gen his. 

fas. 12: 228. (cj ST & 9 ST. C.ayndah, 

Fairmaire, BMNH). NSW, Q. 
viridictps Kerremans, 1898. Ann. Soc. Ent. Belg. 

42: 168 C<3 HT, Australie, Standing, 

BMNH), NSW. 
vi rid /purpurea Carter, 1924. Proc. Linn. Soc. 

NS.W. 4">: 27. (2 6 6 ST. Gcraldton, 

WA. J Clark, AMSA K67294). WA. 
sapphire Carter, 1933. Proc. Linn. Soc. 

N.S.W. 58: 163. (9 HT. Moore R , WA, 

H. W. B.. AMSA K67295)(new syn.). 
watkinsi Barker, 2001 Trans Roy. Soc S. Aim. 

125: 112- (6 HT, Tinonce Rd, Manning 

R.. NSW, S. Watkins, SAMA T 21528). 

NSW. 
wislwoodi (Gory & Laporte, m?i^)(Coraebus). 

Mon, Bupr ii: 15. id ST & 2 9 9 ST, 

MNHN ) SA,V,T 
vr.rna Blackburn, 1891. Trans. R. Soc. S. 

Aust. 14: 299. (HT damaged sex 

unknown. Warburton District, Vicioiia 

liL-m.li collection, NMVA). 
\i intuitu Kerremans. 1898. Ann. Sin. Ent. 

BWjf, 42, 165. (9 HT. Austral., Standing. 

BMNH). 



theryi Kerremans, 1903. in Wytsman Genera 
Ins. fas. 12: 229. (2 6 6 ST & 2 9 9 ST. 
Victoria, Thery, BMNH) 



Discussion 

C. adspersa Kerremans (HT not in BMNH) nab. 

India belongs in another genus. 
C. alberiisi Gestro is not a synonym of C. 

acuducta (Carter 1923) which does not occur 

at the tip of Cape York in the type locality. 
C. comma Gestro (6 ST, MCSNG) = Hypocissei.t 

tatipennis Macleay. 
C. duodecimguttata Gucrin-Meneville. 1830 

(Buprestis). Type species by subsequent 

designation (Duponchel, 1843). 
C fossicollis Kerremans (HT BMNH) ■ Alcinous 

fossicollis (Kerremans). 
C. fulgidicollis Macleay is very uncommon in 

collections and has been confused with other 

species, particularly with C. stellata Barker. 
C. gestroi Kerremans 1892, Mem. Soc Ent. Belg. 

I: 225 is unknown to me; the holotypc is not 

in the BMNH and the species is not listed 

under Ctsseis in Kerremans ( 1903). 
C. maculata Gory &. Laporte was niisidcniifted by 

Carter (1929) who listed C. tusmanka 

Kerremans and C pauperula Kerremans as 

its synonyms; both are good species. 
C. modesta Kerremans was listed as a synonym of 

C. rubicunda Kerremans Carter (1929). I find 

they are separate species, 
C. nigrita Kerremans was listed as a synonym of 

C. notulata (Gennar) by Carter (1929). Carta 

(1923) followed Blackburn's concept of C. 

notulata, which was incorrect. C. mgrita is a 

separate species. 
C, nigrtpennis Macleay was listed by Carter 

(1923) as belonging in Neospades. I find it 

belongs in Cisseis (s.s.). 
C. ohscuru Blackburn is a species that I have been 

unable to Find in any available collection. 
C. oval is Carter could be a synonym of C. opitna 

Thomson. The unique holotypc is a 

discoloured female specimen 
C. rugiceps Thomson belongs in Neospades. 
C. scahiosa (Boisduval) is unknown to me. 
C. setniobscura Kerremans was listed as a 

synonym of C. notulata by Carter (1929). His 

a separate species, much smaller than C 

notulata and common in South Australia 
C. spitota Hope, 1846 (Ann. Mog. Nat. Hist. 17- 

64) (HT. HMOb not seen) is unknown to me. 

Hope indicated that it belonged in anothei 



90 



S. BARKER 



genus. Carter (1929) incorrectly listed it as a 

synonym of C. signaticolUs Hope, 1846; 

spilota has precedence over signaticolUs. 
C. stigmata Gory & Laporte, 1839 selected as type 

species by Bellamy (1998) (invalid). 
C. violacea Kerremans was listed as a synonym of 

C. notulata by Carter (1929). It is a separate, 

smaller species occurring in the vicinity of 

Brisbane, Queensland. 



Acknowledgments 

My thanks for assistance to: C. L. Bellamy, 
Sacramento; M. Moulds, AMSA; T. A. Weir, ANIC; M. 
Kerley, BMNH; A. Samuelson, BPBM; 1. Lansbury, 
HMOE; M. Uhlig, HUMB; R. Poggi, MCSNG; M. 
Hanlon, Sydney; J. Mcnier, MNHN; K. Walker & C. 
McPhee, NMVA; S. Bfly, PMCE; G. B. Monteith, 
QMBA; J. Cools, RMBB; E. G. Matthews, SAMA; J. 
A. Gardner, Waitc Agricultural Research Institute, 
Adelaide. 



References 



BARKER, S. 1998. Selection of leetotypes and 
redescriptions of three Cisseis (Colcoptera: 
Buprestidae) species. Records of the South 
Australian Museum 31: 21-23. 

BARKER, S. 1999a. Designation of a lectotype and 
descriptions of four new species of Australian 
Buprestidae (Coleoplera). Records of the South 
Australian Museum 32: 45-49. 

BARKER, S. 1999b. Designation of leetotypes of three 

species of Cisseis (Coleoptera: Buprestidae). 

Transactions of the Royal Society of South Australia 

123: 155. 
BARKER, S. 2001. Descriptions of twenty one new 

species of Cisseis fsensu stricto) Gory & Laporte 

1839 (Coleoptera: Buprestidae: Agrilinae). 

Transactions of the Royal Society of South Australia 

125: 97-113. 

BARKER, S. & BELLAMY, C. L. 2001. 
Stanwatkinsius, a new genus of Australian jewel 
beetles (Colcoptera: Buprestidae: Agrilinae) with a 



key to known species. Transactions of the Royal 
Society of South Australia 125: 1-14. 

BELLAMY, C. L. 1998. Type species designations in 
the family Buprestidae (Coleoptera). Deutsche 
Entomologische Zeitschrift 45: 9-15. 

CARTER, H. J. 1923. Revision of the Genera Ethon, 
Cisseis and their allies. (Buprestidae.). Proceedings 
of the Linnean Society of N.S.W. 48: 159-176. 

CARTER H. J. 1929. A check list of the Australian 
Buprestidae. Australian Zoologist 5: 265-304. 

DUPONCHEL. P. A. J. 1843 (1841-1848). In 
'Dictionaire universal d'histoire naturelle'. Vols 1-6. 
Ed. C. D. D'Orbigny. Bureau Principal de I'Editeur 
& Renard, Martinet & Cie: Paris. 

GORY, H. L. & LAPORTE DE CASTELNAU, F. L. 
1839. 'Histoire naturelle et iconographie des insectes 
Coleopieres. Monographic des buprestides". Volume 
2, livraisons 25-35. P. Dumenil: Paris. 

KERREMANS C. 1903. Genera Insectorum 12: 1-338. 



THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS 
COMMITTEE 10 MARCH 1921 - 30 OCTOBER 2001 

Wolfgang Zeidler 



Summary 

The Flora and Fauna of South Australia Handbooks Committee has been operating for 80 years 
producing a series of handbooks on the major elements of the South Australian natural 
environment. The main role of the Committee was to encourage and persuade specialists to write 
books on the flora and fauna for no financial reward. The completed manuscripts were then edited 
by members of the Committee and, in the past, were printed by State Print or its equivalent and 
published by the State Government Printer at the State Government's expense. Several years ago 
this part of the Government' s operations ceased, with no provision made for the further printing of 
handbooks. 



THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS COMMITTEE 
10 MARCH 1921 - 30 OCTOBER 2001 



WOLFGANG ZEIDLER 



ZEIDLER, W. 5002 The Flora and Fauna of South Australia Handbooks Committee 10 March 
1921 - 30 October 2081, Kecords of I he Saulh Australian Museum 35(1) 91 -95 

Wolfgang Zeidlcr, South Australian Museum, North Tetrace, Adelaide, South Australia 5000; 
and Visiting Research Fellow, Adelaide University. Adelaide, South Australia 5005. 
Manuscript received 13 December 2001, 



The Flora and Fauna of South Australia 
Handbooks Committee has heen operating lor 80 
years producing a series of handbooks on the 
major elements of the South Australian natural 
environment The main role of the Committee was 
to encourage and persuade specialists to write 
books on the flora and fauna for no financial 
reward. The completed manuscripts were then 
edited by memhets of the Committee and, in the 
past, were printed by State Print or its equivalent 
and published by the Slate Government Printer at 
the State Government's expense- Several years 
ago this part of the Government's operations 
ceased, with no provision made for the further 
printing of handbooks. 

Since the privatisation of the Government Printer 
the Committee has sought other means to publish 
its books and flip last two handbooks have heen 
printed with the help of the Board of the Botanic 
Gardens ('Fungi') and the SA Research & 
Development Institute (Aquatic Sciences) ("Marine 
Invertebrates HI'). However, it became very clear 
to the Committee that the only way it could 
continue was to establish a publications fund 

Initially, the Stale Government was approached 
because of its traditional .support for the 
Committee, but this proved fruitless, although the 
Department of Environment & Heritage and the 
South Australian Museum did offer some support 
if funds could also be found from other sources. 
Various philanthropic trusts were approached and. 
although some were sympathetic, no funding was 
forthcoming. Similarly, all three of the State's 
universities, whose students and staff are amongst 
the main users of the handbooks, declined to assist. 
The formation of B partnership with a publisher 
was also considered, but rejected as it would 
potentially compromise the original intention of the 
Committee because handbooks with commercial 
potential would be favoured. 



Left with no alternative, the Handbooks 
Committee Teluetantly concluded that it could not 
continue to function as originally intended and 
should be disbanded. The last meeting of the Flora 
and Fauna of South Australia Handbooks 
Committee was accordingly held On 30 October 
2001. 

With the demise of the Handbooks Committee, 
which was once the envy of all other states, it is 
important to record its history and major 
achievements. 

The Committee began in 1921 when Prof E. 
Wood-Jones. Prof T. B. G. Osboni and Prof J. B. 
Cleland presented the idea to the then Premier Mr 
H. Barwell, The concept was for authoritative 
handbooks to be written by experts in their field 
without remuneration, to be published by the 
Government Printer at its convenience, and to be 
sold at a low price so as to be affordable to the 
wider community. Here follows an extract 
recording the foundation of the handbooks series, 
prefacing Eiehler's (1965) 'Supplement to J. M 
Black's Flora of South Australia'. 

After the first World War. the South Australian 
Branch of the British Science Guild was- very 
active in devising ways for increasing scientific 
knowledge and its applications At the time an 
up-to-date account of die Flora of our Slate was 
a pressing need. Willi this view, Professor J. 8 
Cleland suggested that die Science Guild might 
undertake to arrange Tor the preparation of a 
scries tit Handbooks dealing wilh (he Fauna and 
Flora of South Australia. If these were- prepared 
in an honorary capacity by leading authorities, 
the Government might well be asked to publish 
ihem, as it were, on a pound for pound basis 
The Brunch accepted the suggestion and it was 
arranged that a deputation of Professors P. 
Wood Jones, T. G. B. Osborn and J. B. Cleland 
should wait on the Hon. The Premier Mr., 
afterwards Sir Henry Barwell, wilh the proposal 



92 



W. ZEIDLER 



Fortunately Professor Cleland had prepared a 
memorandum on the subject in the following 
terms and this document was handed to the 
Premier on 25* February, 1921. 

Proposed Scheme For The Preparation Of 

Handbooks On The Fauna And Flora 

Of South Australia 

I. There is an almost complete lack in South 
Australia of any handbooks dealing with 
the Fauna and Flora of the State. 

II. In consequence, many individuals with 
scientific tastes are unable to develop 
these. This is a great loss to the State, as 
these persons, with suitable handbooks 
available, might be led on to contribute 
voluntarily to the State additions to 
scientific knowledge of great value to our 
pastoral, agricultural and other interests. 

III. A scheme under the auspices of the S.A. 
Branch of the British Science Guild is 
proposed for the preparation of a series 
of such handbooks to be edited by a sub- 
committee of experienced workers, the 
individual parts to be prepared by 
specialists (in an honorary capacity). 

IX. The scheme would be spread over a 
period of 10-15 years. 

X. Parts would be issued as completed by 
the authors. None could be available 
during this financial year. One or two 
may be so for 1921-1922. 

XI. The work involved will be highly skilled 
and laborious. In the interests of science 
the authors will be prepared, we believe, 
to devote their time and knowledge to this 
work without fee, if a means of 
publication can be obtained. 

XII. The value to the community of the works 
of these authors, thus offered as a gift to 
the State and world of science, can be put 
at a very high figure. Would the 
Government be prepared to accept this 
offer of gratuitous service and as a return 
- on the "pound for pound subsidy" 
principle - arrange for the publication of 
the Handbooks by the Government 
Printer? 

XIII. As the work will be spread over many 
years, this should not necessitate any 
additions to the staff of the Government 
Printer, the printing of the parts being 
proceeded with by him as opportunity 
offered. The actual out-of-pocket expense 
to the State would consist, under these 
circumstances, of the paper only. The 
parts as completed might be submitted to 



the Premier or to the Minister of 
Education, and laid on the table of the 
House and ordered to be printed and thus 
be deall with as Parliamentary Papers. 

IX. Many of the parts will require to be 
illustrated. In many cases, blocks are 
already prepared which doubtless could 
be used and so reduce expense. 

X. The educational value of these handbooks 
will be great. A copy of each might be 
placed with advantage in every public 
school in the State, when it could be made 
available not only to teachers and 
scholars but to the public of the district 
as well. To meet this demand and to 
enable distribution to be made to private 
individuals (at a nominal price to induce 
the wide use of these handbooks) about 
1 ,500 copies of each should be printed. 

XI. Probably from 20 to 30 handbooks would 
eventually be published (over the period 
of 10-15 years), the sizes varying from 
about 20 to 150 pages according to 
subject matter. 

Professor CLELAND, Adelaide University. 
Professor WOOD JONES, Adelaide University. 

The Premier read through the memorandum and 
said, "A very generous offer, gentlemen. I shall 
lay it before Cabinet." Professor Cleland 
received from the secretary to the Premier, a 
letter dated 10* March, 1921 in the following 
terms accepting the offer. 

"With reference to the proposed scheme under 
the auspices of the South Australian Branch of 
the British Science Guild for the preparation of 
a series of handbooks on the Fauna and Flora of 
South Australia which was submitted by 
yourself and Professor Wood Jones to the 
Premier on the 25* ultimo, I am directed by the 
Premier to inform you that in consideration of 
the contributors in the compilation of the 
handbooks undertaking the work in an honorary 
capacity the Government is prepared to 
undertake the printing of the publication at the 
Government Printing Office at the expense of 
the State." 

In recent years the Chief Secretary of South 
Australia decided that the Government Printer 
should publish the handbooks on a commercial 
basis, rather than that the Handbooks Committee 
should apply each year for a government grant 
for publication. Consequently there is now no 
delay in printing when the authors offer their 
completed manuscripts. 

The British Science Guild has long since ceased 
to exist but the work of the Handbooks Committee 
survived until now. The main aim of the 



THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS COMMITTEE 



93 



Committee has always been to produce books that 
document, in a rigorously scientific way, the 
biological heritage of the State, although in recent 
years the Committee encouraged authors lo 
broaden their focus to southern Australia in 
general. Perhaps the most outstanding work 
published by the Committee is the 'Flora of South 
Australia". The remarkable J. M. Black produced 
the four- volume work single-handedly and revised 
the first three volumes, working until he was 96. 
Revision of the fourth volume was completed 
after his death. For South Australia's 
scsquicentenary a completely revised and 
rewritten new edition was paxluced, making this 
State the only one at the time to have a 
comprehensive up-to-date record of State flora. 

There are also other monumental works for 
which other states have no equivalent (see list of 
handbooks) and. while sunie volumes are out of 
dale, many have been updated or reprinted to meet 
the demand of scientists, naturalists and students. 
Ii seems remarkable thai Hale's 'The Crustaceans 
Of South Australia' (1927, 1929), reprinted in 
1976, is still used as a basic text, although very 
much out Of date and only providing a limited 
coverage of the group. 

It is clear that the handbooks provide invaluable 
data for research workers, not only in South 
Australia, or even nationally, but internationally 
as well. Indeed, the handbooks series has been the 
envy of scientists in other states. Their 
contribution to secondary and tertiary education, 
as significant textbooks, is well recognised. For 
example, the four-part 'Flora', 'Fishes' and the 
three-part 'Marine Invertebrates of Southern 
Australia' series are essential texts for tertiary 
students. Unfortunately, the later volumes 
produced by the Government Printer were 
managed on a commercial basis and were 
remaindered within two years, so that they were 
only available for a relatively short period- 
Future plans for new editions of 'Fishes', 
Amphibians', 'The Vegetation of South Australia'. 
1'iustacea' (as pan of the 'Marine Invertebrates' 
series) and 'Flora' have been abandoned, although 
llicy may be taken up by others. In particular, the 
State Herbarium is in the process of producing an 
electronic version of the 'Flora', and the South 
Australian Research & Development Institute 
(Ai|ii;uic Sciences) intends to proceed wilh future 
vMiuincs of the 'Marine Invertebrates' series as well 
as revising earlier volumes. 

As part of the winding up process, the 
Committee has transferred copyright of the 
botanical tests lo ihe Board of the Botanic Gardens 



and Stale Herbarium, and of the zoological and 
ecological texts to the Board of the South 
Australian Museum. Copies of correspondence and 
minutes of the Handbooks Committee have been 
deposited with State Archives. 

Titles Issued By The Flora And Fauna Of- South 
Australia Handbooks Committee 

1922 'Flora of SouLh Australia'. Part 1 
Cyathaceae- Orchidaceae. J. M. Black- 
Second edition 1943. Reprinted 1948. 
Reprinted (facsimile) I960, 1972. Third 
edition, Lycopodiaceae - Orchidaceae 
(revised by J. P. Jessop) 1978. 

1923 'The Fishes of South Australia'. E. R. 
Waite. 

Reprinted 1962. 

1923 "The Mammals of South Australia'. Part I. 
The Monotremes and the Carnivoro'j- 
Marsupials. F. Wood-Jones- 
Reprinted in one volume with parts II 
(1924) and III (1925), with an Introduction 
by P. Crowcroft and a Selected 
Bibliography by J. H. Calaby 1968. 

1924 'Flora of South Australia'. Pari II 
Casuarinaceae - Euphorbiaceae. i. M 
Black. 

Second edition 1948. Reprinted 1963. 
Reprinted (facsimile) 1977. 

1924 'The Mammals of South Australia'. Part 
II. The Bandicoots and the Herbivorous 
Marsupials. F. Wood-Jones. 
Reprinted in one volume wiih parts 1 
(1923) and III (1925), as above, 1968. 

1925 'The Building of Australia and the 
Succession of Life' with special reference 
to South Australia. Part 1. W. Howchin, 

1925 'The Mammals of South Australia'. P.m 
III The Monodclphia. K Wood-Jones. 
Reprinted in one volume with parts I 
( 1923) and U (1924), as above, 1968. 

1926 Flora of South Australia' Part 111 
Meliaceae — Scropiiulariaceac. J. M. 
Black. 

Second edition, Callitrichaccac - 
Plumbagmaceae 1952. Reprinted I9M. 
Reprinted (facsimile) 1977. 

1 927 'The Crustaceans of South Australia'. Part 
1. H. M. Hale. 

Reprinted in one volume with part II 
(1929)1976. 



94 



W ZEIDLER 



1928 'The Building of Australia and the 
Succession of Life' with special reference 
to South Australia. Part II. Mcsozoic and 
Cainoz.oic W. Howchin. 

1929 'Flora of South Australia'. Part TV. 
Bignoniaccac - Compositae. J. M. Black. 
Second edition, Oleaceae - Compositae 
(revised by E. L. Robertson) 1957. 
Reprinted (facsimile) 1980. 

1929 'The Crustaceans of South Australia' Part 
n. H M. Hale 

Reprinted in one volume with part I (1927) 
1976. 

1929 'The Reptiles and Amphibians of South 
Australia'. E. R. Wake. 

1930 'The Building of Australia and Lhc 
Succession of Life' with special reference 
10 South Australia Part III. Pleistocene. 
W. flowchin. 

1934 "Toadstools and Mushrooms and other 
Larger Fungi of South Australia,' General 
introduction; Toadstools and Mushrooms. 
J. B. Cleland. 

Reprinted in one volume with pari It 
(1935), including 'Notes on some Edible 
and Poisonous Fungi' by P. H. B. Talhoi 
1976 

1935 'Toadstools and Mushrooms and other 
Larger Fungi of South Australia". 
Polypores, Coral Fungi and remaining 
Hymenomyeetcs; Puff-balls, Jelly-like 
fungi, the larger Ascomycetes and 
Myxomycetes. J. B. Cleland. 
Reprinted in one volume with part 1 
(1934), as above. 

1936 'The Seaweeds of South Australia'. Part I. 
Introduction and The Green and Brown 
Seaweeds. A. H. S. Lucas. 

1937 The Vegetation of South Australia'. J, G. 
Wood 

1938 'The Molluscs of South Australia'. Part I. 
The Pelecypoda. B. C. Cotton & F. K. 
Godfrey. 

1939 'Primitive Insects of South Australia'. 
Silverfish, Spnngtails and their allies. H. 
Womersley. 

1 940 'The Molluscs of South Australia'. Part II. 
Seaphopoda, Cephalopoda, Aplacophora 
and Crepipoda. B. C. Cotton & F. K. 
Godfrey. 

1947 'The Seaweeds of South Australia'. Part 



1959 
1961 
1962 
1964 
1965 

1966 

1972 
1974 

1 975 

1976 

1978 
1978 
1979 
1980 

1980 
1982 

1984 
1984 
1986 



II. The Red Seaweeds. A. H. S. Lucas & P 
Perrin. 

'South Australian Mollusca'. 

Archaeogastropoda. B. C, Cotton. 

'South Australian Mollusca", Pelecvpoda. 

B. C. Cotton. 

The Marine and Freshwater Fishes of 
South Australia'. T D. Scott. 

South Australian Mollusca'. Chitons. B. 

C. Cotton. 

'Supplement to I. M. Black's Flora of 
South Australia' (second edition 1943- 
1957). H.J. Eichler. 

'Aboriginal Man in South and Central 
Australia'. Part I. Edited by B. C, Cotton. 

Only one part was published. 

'The Vegetation of South Australia' R L. 
Specht. Second edition. 

'The Marine and Freshwater Fishes of 
South Australia' T D. Scott, C. J. M. 
Glover & R. V. Southcott. Second edition. 
Reprinted (facsimile, wilh 16 pp of colour 
plates) 1980. 

'Plant Feeding and other Bugs (Hemiptcra) 
of South Australia'. Helcroplera Part J. G. 
F. Gross, 

'Plant Feeding and other Bugs (Hemiptera) 
of South Australia". Heteroptera Part n. G. 
F. Gross, 

'Butterflies of South Australia'. R. H. 
Fisher. 

'Amphibians of South Australia', M. J. 
Tyler. 

'Lichens of South Australia'. R. B. Filson 
& R. W. Rogers. 

South Australia' 



'Mosses of 
Catchcside. 



D. G 



D. J. E. 



'Acacias of South Australia' 
Whiblcy. 

'Marine Invertebrates of Southern 
Australia'. Part I. Edited by S. A, 
Shepherd & 1. M. Thomas. 

'Psylloidea of South Australia'. F. D. 
Morgan. 

'The Marine Benthic FJora of Southern 
Australia'. Part I. H. B. S. Womersley 
'The Dynamic Partnership: Birds and 
Plants in Southern Australia' Edited by H. 
A. Ford & D. Paton. 



THE FLORA AND FAUNA OF SOUTH AUSTRALIA HANDBOOKS COMMITTEE 



95 



1986 'Flora of South Australia. Volumes I-IV. 
Edited by J. P. Jessop & H. R. Toelken. 
Fourth edition (third edition consisted of 
part I only). 

1986 'The Ecology of Forests and Woodlands 
of South Australia'. Edited by H. R. 
Wallace. 

1987 'The Marine Benthic Flora of Southern 
Australia'. Part II. H. B. S. Womersley. 

1989 'Marine Invertebrates of Southern 
Australia'. Part II. Edited by S. A. 
Shepherd & I. M. Thomas. 



1990 'Orchids of South Australia'. R. Bates & 
J. Z. Weber. 

1992 'Acacias of South Australia'. D. J. E. 

Whibley & D. E. Symon. Second edition. 
1994 'The Fishes of Australia's South Coast'. 

Edited by M. F. Gomon, J. C. M. Glover 

& R. Kuiter. 

1997 'Larger Fungi of South Australia'. C. A. 
Grgurinovic. 

1997 'Marine Invertebrates of Southern 
Australia'. Part III. Edited by S. A. 
Shepherd & M. Davies. 



Dr Wolfgang Zeidler (Hon. Secretary), on behalf of the Flora and Fauna of South Australia Handbooks Committee 
— Em Prof William D. Williams (Chairman), Dr Sue Barker (Botanical Editor), Dr Margaret Davies (Zoological 
Editor), Dr Shelley Barker, Ms Robyn Barker, Dr John G. Conran, Mr Peter Copley and Dr Brian D. Morley. 



OBITUARY GRAEME LLOYD PRETTY 1940 - 2000 

Helen Tolcher 

Summary 

Graeme Lloyd Pretty was born in Melbourne on 25 June 1940. His father was in the RAAF and the 
family soon moved to Richmond NSW, where Graeme spent his childhood and went to school. 
After graduating from Sydney University BA (Hons) in Classical Archaeology and Dip Ed, he 
contemplated undertaking a PhD with Jack Golson at the Australian National University. However, 
he began working at the South Australian Museum in January 1962 as Assistant Curator of 
Anthropology under Norman Tindale, from whom he learned much and with whom he kept in touch 
long after the latter' s retirement. His first interest was in Melanesian prehistory - he might well 
have made this his major field of work. 



OBITUARY 
GRAEME LLOYD PRETTY 

1940-2000 




Graeme Prelty and Henry Pollach (left) at Roonka, 8 August 1971. 
Roonka photos R83r 2. South Australian Museum 



Graeme Lloyd Prctly was bom in Melbourne on 
25 June 1940. His father was in the RAAF and 
the family soon moved to Richmond NSW, where 
Graeme spent his childhood and went to school. 
After graduating from Sydney University with a 
BA (Hons) in Classical Archaeology and Dip Ed, 
he contemplated undertaking a PhD with Jack 
Golson at the Australian National University. 
However, he began working at the South 
Australian Museum in January 1962 as Assistant 
Curator of Anthropology under Norman Tindale, 
from whom lie learned much and with whom he 
kept in touch long after the latter*s retirement. His 
first interest was in Melanesian prehistory — he 
might well have made this his major field of work. 

In 1964 he was appointed Assistant Curator 
of Archaeology, and archaeological fieldwork 
was added to his duties. He excavated or 
assisted at a number of sites on the lower River 
Murray, seeking the advice of John Mulvaney 
who had recently excavated Fromnt's landing. 
At the same time he continued research into 



Melanesian culture, including six weeks 
fieldwork in the Southern Highlands of Papua 
New Guinea (PNG) in 1968-69. Subsequently, 
he was invited to review the functions of the 
PNG Museum and An Gallery. Later, he also 
advised the Australian National Gallery on its 
Melanesian collections. 

In 1968 Prelty began a project that was to 
become the most important one of his working 
life. A salvage operation to remove a surface 
campsite and scattered human remains from a 
sandhill beside the River Murray at Roonka. near 
Blanchetown, developed into a major excavation 
that would attract worldwide interest. At that time 
he was unique in having consulted with the 
relevant Aboriginal group and obtained approval 
before beginning work — he continued this contact 
throughout and after the conclusion of the dig. 
This rapport with the Aboriginal people 
undoubtedly contributed to making possible the 
excavation on the scale and significance that was 
achieved. 



98 



H. TOLCHER. J. PRESCOTT & 3. MULVANEY 



Despite the disadvantages and difficulties of 
working on a surface fully exposed to gale-force 
winds and summer heat rising to the SCTC range, 
he retrieved a wealth of archaeological material 
and information from a site which proved to have 
been occupied for some IK 000 years In all, 216 
complete or fragmentary burials were uncovered. 
The grave goods were unprecedented at that lime 
In addition td almost every variant of human 
mortuary practice, it also gave evidence for 
hitherto undocumented customs, of which there 
were well over 150 examples, 

Roonka looms large for its deep significance 
Here is a statistical sample from one place, across 
some thousands of years, permuting health, injury 
Bnd dietary studies of the population Pretty 
correctly saw that aspects of the rituals involved — 
the mode of bunal, including grave goods, and 
the antiquity of the rituals — presented a major 
window to past belief and behavioural systems. 
Nu other burial ground has produced no much 
evidence. 

The almost exclusively amateur workforce of 
volunteers became, over the eight years of active 
ficUtwork, a team of patient, dedicated, highly 
skilled excavators Under Pretty's direction, the 
site was stnctly controlled, meticulously recorded 
and kept secure from outside interference. The 
standard and scope of the work attracted 
substantial financial support aA well as visitors 
from abroad, including osteo-arehaeologists and a 
delegation from the People's Republic of China. 
One group will remember the visit not only for 
the archaeology but because their bus became 
bogged to the axles on the Roonka Flat. 

Pretty was appointed to the position of Senior 
Curator of Anthropology and Archaeology in 
1975. 

The section of Pretty's work that has been best 
documented is probably that dealing wiih 
chronology. He had a critical appreciation of the 
role of physical dating methods in archaeology 
and sought to create a temporal framework for the 
finds at Roonka. The chronology of the site was 
determined by a larger number of different dating 
methods than any other archaeological site in 
Australia, including tlicrmoluminesccnce, 
palaeomagnetism. uranium fluorimctry and C-14 
dating, both conventional and AMS. 

ft is not generally known ibai he was 
responsible for planting the idea for developing 
luminescence dating in the Physics Department at 
the University uf Adelaide in convcrsalion with 
Professor John Prcscoll. This was in 1972, at the 
very infancy of the thermoluminescence 



technique. The earliest Adelaide work on 
Uiennoluminescence dating per sc was based on 
samples from Roonka. 

Pretty extended his ficUtwork research to Island 
Melanesia in 1971 and 1973, finding additional 
Support from sources outside Australia, However, 
the demands of attempting to elucidate the 
intricacies of the Roonka information, together 
with consiraints on finance and staffing, resulted 
in the Melanesian work being put aside — apart 
from ongoing work, under his direction, on the 
sorting of the Foreign Ethnology reserve 
collections and their eventual rehousing at Kent 
Town in 1986. As Senior Curator of Archaeology, 
Pretty's work was directed almost exclusively to 
organising the mass of information from the 
Roonka site in order to arrive at what he 
considered an adequate assessment on which to 
base his final report. During the 1980s a master 
index of the Roonka data was prepared. 

The Third Australasian Archaeometry 
Conference was held in Adelaide in 1988. Pretty 
devised the theme of the Symposium, 'Early Man 
in the Southern Hemisphere', which was designed 
io give a specifically archaeological balance to the 
ptUgtam He arranged invasions and finance for 
the international contributors; G Delibrias, G. J. 
Bartstraa and H f. Deacon; and organised ihe 
conference excursion and wiotc the field nolo. 

In his own contribution to the symposium, a 
paper analysing the cultural chronology of 
Roonka. it is possible tc see a foretaste of his 
overall interpretation of Roonka in its wider 
context. An early indication is also found in his 
exciting and innovative 1977 paper, 'The cultural 
chronology of Roonka Flat'. In it he remarked 
'This present paper therefore celebrates the 
author's satisfaction that the archaeology is now 
sufficiently clear lo commit preliminary notice ol 
it to paper' Yet, during the following 23 years, tic 
produced no paper with the promised detail. It ts 
difficult tor those of us who knew him to avoid 
reflection on his apparent reluctance to put his 
ideas on paper where they could be exposed to the 
critical appraisal of his peers. He did publish 
sections of the work with his collaborators, but 
the overall synthesis was yet to come. Perhaps he 
was sensitive to possible criticism, perhaps the 
very volume of his material was an obstacle. It 
seems likely that he laboured over the precise 
language with which to present a polished linai 
monograph. 

In 1982 restructuring of the South Australian 
Museum's divisions and staffing made Pretty one 
of a number of Curators of Anlhropology. He 



OBITUARY - GRAEME LLOYD PRETTY 



accepted a severance package in 1994, continuing 
his woik on the Roonka report until 1996, when 
he was stricken with viral encephalitis, front 
which he never fully recovered. He died on 
6 November 2000. 

In his work Pretty was patient, a seeker of 
precision in language that was baffling to many, 
and single-minded to a degree thai ignored 
workplace relationships and, ai times, made it 
difficult for people to work with hirn. His memory 
was exceptional. The breadth of his reading was 
remarkable and was reflected in his large personal 
library, He left the Museum's Foreign Ethnology 
collection well organised and totally accessible, 
and the Roonka material fully recorded and 
stored. He gained the respect of his field workers, 
instructing them in aspects of Ahorigina! culture 
and inspiring in them a dedication and loyally to 
the project that will ensure the publication of the 
final report. 

Graeme married Dana in 1973 and they had one 
daughter. To those who knew him well he was a 
kind and considerate friend. At home he was a 
gracious and amusing host, an erudite and 
entertaining conversationalist, a devoted husband 
and a proud and loving father. 



Authored And Collaborative Work To 1 995, 
Listed By Field 

Archaeology 

PRETTY, G. L. 19b4. Stone objects excavated in New 
Guinea. Man 64, 138: 17. 

PRETTY. C. L. 1964, A stone figure of a crested 
cockatoo from Melanesia. Man 64. 220: 183-184. 

PRETTY. G. L. 1965. Two stone pestles from Western 
Papua and their relationship to prehistoric pestles and 
mortars from New Guinea. Records of the South 
Australian Museum 15. I: 120-130. 

PRETTY, G. L. (wiih M. J. TYLERl. 1966. A bizarre 
refuge for some Austtalian Icptodactylid frogs, 
Remtflt of the Snath Australian Museum 15. 2; 360- 
361, 

PRETTY. G, L 1967 Excavations at die Anglo-Suxun 
site of Wallifigford. Berksfntc. England. 1966 
.luiirrtfil of the Anthrof.-ological Society of Smith 
Austtalta 5, 2; 3-5. 

PRETTY. G. L. 1967 Rescue excavation of an 
aboriginal grave. Tailcm Bend. South Australian 
Naturalist 41. 4 9-11. 

TTY, G. L. (with S. A. GALLUS). 1967. The 
anthropology and archaeology of die Nullatbor Plain. 
South Australia. Pp. 47-49 in 'Caves of the 
Nullarbor'. Eds. J R. DUNKLEY & T M. L. 
WIG LEY Addiii.k' 



PRETTY, G L, 1968. Excavation of Aboriginal graves 
at Gidgealpa, South Australia. Records of the South 
Australian Museum 15, 4: 671-677. 

PRETTY. G. L 1968 Two more prehistoric stone 
artifacts from Western Papua. Records of the Soutli 
Australian Museum 15, 4: 693-697 

PRETTY. G. L, 1969. Prehistoric background to 
Australia. Tradition: Journal of the History 
Teachers' Association ISA.) May. Pp 13-23. 

PRETTY. G. L. 1969. Excavation of an Aboriginal 
cemetery near Blanchctown. River Murray, Soudi 
Australia. Paper read to die 4 1 si Congress of die 
Australian and New Zealand Association for the 
Advancement of Science, Adelaide, August 1969. 15 
pp., 10 illus. 

PRETTY, G. L. 1970. Excavation of an Aboriginal 
cemetery on Roonka Station, River Murray. Kalori, 
Journal of the Museums Association of Australia 38: 
17. 

PRETTY. G. L. 1971. Excavations at Roonka Station. 
Lower River Murray, South Australia, 1968-1970 
Journal of the Anthropological Society of South 
Australia 19. 9: 6-15. 

PRETTY. G. L. 1971. Further investigations into 
Melanesian culture-history; South Australian 
Museum field research in Island Melanesia, 1971. 
Kalori: Journal of the Museums Association of 
Australia 42: 89-95, 

PRETTY, G. L. 1972. "Report of an Inspection of 
Archaeological Sites and Field Monuments in the 
Territory of Pupua New Guinea'. Papua New Guinea 
Public Museum and An Gallery: Port Moresby, vii + 
50 pp., illus. 

PRETTY, G. L. 1975. Archaeology in South Australia: 
a report on recent work. Australian Archaeology: 
Australian Archaeological Association Newsletter 3: 
32-39. 

PRETTY. G. L. 1975, 'Social Change in Prehistoric 
Australia; the Evidence from Roonka'. Paper read to 
the 46th Congress of the Australian and New Zealand 
Association for ihe Advancement of Science. 
Canberra. 12 pp.. 16 illus 

PRETTY, G L. 1976. The excavations it Roonka Flat, 
South Australia: an insight into ancient Australian 
society. Actes el Proc IX Cong res, Union 
Internationale des Sciences Prehistnriques el 
Proiohisloriques, Nice. Section 6. 2. 100-112. 

PRETTY, G. L. W7. Atchaeology m South Australu. 
Pp 40-54 in South Australian Yearbook' 
Australian Bureau of Statistics: Adelaide. 

PRETTY. G. L. 1977. The cultural chronology of" the 
Roonka Flat: a preliminary consideration. Pp. 28Si 
331 in 'Stone Tools as Cultural Markers. Change. 
Evolution, Complexity" Ed. R. V. S. WRIGHT 
Australian Institute Of Aboriginal Studies: 
Canberra. 

PRETTY, G. L, 1978. Review of V, D. Watson & J. D. 



100 



H. TOLCHER, J. F'RESCOTT & J MULVANEY 



Cole, 'Prehistory of the Eastern Highlands o(" New 
Guinea'. The Artefact 5: 98-102. 

PRETTY, G. L. 1980. Prehistory. Pp. 9-17 in 
'Proceedings of the Fiftieth Anniversary Seminar of 
the Anthropological Society of South Australia'- 
Adelaidc. 

PRETTY, G. L. 1981 Trial excavation of an Aboriginal 
camp site and site survey. Kinehega National Park, 
Pp. 53-63 in 'Darling Surveys, V. Ed, J. M. HOPE. 
Occasional Papers in Prehistory 3. Australian 
National University Canberra. 

PRETTY, 0. L 1982. 'The Archaeology of Roonka. 
Examining the Human and Environmental History of 
an Australian Ptace'. Paper read to the Nature 
Conservation Society of South Australia, Adelaide. 
23 pp.20 illus. 

PRETTY G L. 1982. Prehistory of the Australians: the 
view from Roonka. University of Adelaide, 
Department of Classics, Lecture Seiies-Archaeologv 
IH: 1-11. 

PRETTY. G. L. (with R. C. PATON St. R. D. J. 
WEATHERSBEB). 1983. Ttibal man. Pp. 115-126 
in 'Natural history of 'lie South East of South 
Australia'. Eds. M J TYLER, C. R. TWIDALE, J 
K. LING & J. W HOLMES Royal Society of South 
Australia: Adelaide. 

PRETTY, G. L I with T. J. GARA). 1983, Scientific 
background, in 'Roonka Conservation Park 
Management Plan'. Ed. P J. HUGHES, ANUTECH 
Services- Canberra 

PRETTY, G, L. 1984. Review of John Aubrey (cd.), 
"Monumenta Britannica or a Miscellany of British 
Antiquities (parts 1 & 2)'. Mankind Id. 3: 254-6. 

PRETTY, G. L. 1986 The prehistory or South 
Australia Pp. 3-62 in 'The Flinders History of South 
Australia: Social Histoiy. 3'. Ed. E RICHARDS 
Wakefield Press: Adelaide. 

PRETTY. C. L. (with R. S. MERRILLEES & C A 
HOPE). 1990. 'Living with Egypt's Past in 
Australia'. Museum of Victoria. Melbourne, viii + 
78 pp., illus 

PRETTY. G. L, 1990. The significance or the tombs of 
evolved prehistoric hunters and gatherers; Southern 
Australia and Southern France compared. (Invited 
address, Royal Society of South Australia. May. 
1990.) Newsletter. Royal Society of South Australia, 
June Pp. 4-7. 



The following are in manuscript form and it is 
intended thai they will be published in due course. 
PRETTY. G. L. 'Archaeological Investigations at 
Roonka, Part I - Description of the evidence gathered 
from the Karmakalingbila Ngaiawang territory: 
Geological and geoehronologicai background; 
cultural landscape and stratification; ecological, 
demographic and cultural finds'. Text, labies, illus. 



PRETTY, G, L. (with G K. WARD) (eds). 
'Archaeological Invesligaliuns at Roonka, Part 2 
Ethnoanalogical arid compaiativc analysis: 
Conjectures about territory, society and ideas in the 
Karmakalingbila Ngaiawang territory in the light of 
its Murray Darling Basin context'. (Accepted for 
publication in British Archaeological Reports: 
International Scries.) 

PRETTY, G. L. 'Archaeological Investigations at 
Roonka, Part 3 - Ethnodeduciive and systemic 
analysis* Conjectures ahout the mechanics and 
dynamics of cultural change in the 
Karmakalingbila Ngaiawang territory in the light 
of its Central Southern Australian context'. Text, 
tables, illus. 

PRETTY, G. L. 'Archaeological Investigations at 
Roonka. Part 4 - Decoding the symbol inventory of 
the Karmakalingbila Ngaiawang sequence: Rcstonng 
immediate intelligibility and ultimate causal identity 
to culturally unfamiliar modes or action through 
appeals to phyletic behavioural repertoires'. Text, 
labies, illus. 

PRETTY. G. L. 'Archaeological Investigations at 
Roonku, Part 5 • The explanation of prchistm it 
hunrer-gatherer culture change in the 
Karmakalingbila Ngaiawang territory during the 
Holocene, tests of confirmation and concluding 
synthesis* Text, tables, illus. 



Art 

PRETTY, G. L 198$. A turtle shell mask of Torre* 

Strait type in the Maclcay Museum, University of 

Sydney. Records of the South Australian Museum 

-4. 3 .421-42-5. 
PRETTY, G L. 1971 The National Collection <if 

Primitive Art, Canberra Art and Australia 8, 4: 324- 

329. 

PRETTY, C. L. 1971. Common on R McOhcc. 

'Differential artistic productivity in the Eskimo 

cultural tradition'. Current Anthropology 17. 2: 216. 
PRETTY, G. L 197IH. 'Arts Of Melanesia' Anvinnli- 

Festival (jf Arts !97tV ("ixhibttion Catalogue,) 1 ).i\ -ni 

Jones Gallery: Adelaide. 



Cultural nuritase and public policy 

PRETTY. G L. 1970. 'Report on the Commonwealth 
Collection of Primitive An from New Guinea and the 
Pacific Islands'. Commonwealth Art Advisory 
Board: Canberra, iv + 78 pp., illus. 

PRETTY. G. L. 1971. Museums and controls on the 
e>port of cultural property Kalori: Journal of the 
Mti.\eums As.\oi iatioti of Australia 42: 89-95 

PRETTY. G L (General Editor) 1977 'National Folk 
Province Feasibility Study Papers'. Commonwealth 
Department lor the Environment, Housing and 



OBITUARY -GRAEME. LLOYD PRETTY 



101 



Cornmunily Development and South Australian 
Department for the Environment: Adelaide, as 
follows: 
No. I PRETTY, G L. -Tire Ngaiawang Folk Province. 
A Proposal' 

No. 2 PRETTY. G. L, 'The National Folk Province 
Concept - Specification lor a feasibility study'. 

No. 3 ROWNEY, B. G. 'Kapunda - Historical 
development of its town plan'. 

No. 4 PRETTY. G. L. 'The National Folk Province 

Concept and the National Estate' 
No 5 KALIBATAS. E. 'Craigie's Plains: Construction 

and materials analysis of a historic vernacular sheep 

station' 
No. 8 SPIERS, G. K 'Survey of Heritage Agencies in 

South Australia', 
No 9 ROWNEY, B. G 'Surviving cottages in Mine 

Square. Kapunda'. 

No 10 PRETTY.. G. L. (with B. C. ROWNEY & P. F 
DONOVAN) 'Ngaiawang Folk Province: 
Preliminary inventory of ihe cultural lundscape'- 

No. II SPIERS, G. K. 'The administration of heritage 
properties through Local Government: I, The South 
Australian Planning and Development Act 1996- 

1975 . 

No. 12 DONOVAN, P. F. & SCHEPPERS. K H. 
'Report on the Kapunda Colonial Festival, 1976'. 

No. 13 SPIERS. G. K 'The administration of heritage 
properties through Local Government: It, The 
Municipal Heritage Trust', 

No 15 DONOVAN, P. F. Woakwine Folk Province: 
Outline sketch of cultural landscape'. 

No 16 DONOVAN, P. P & SCHEPPERS. K. H. "The 
administration of heritage property through Local 
Government: III, Report from the Heritage Officer. 
District Council of Kapunda'. 

No 19 ROWNEY, B. C. & DONOVAN, P, F. 'Old 
Legislative Council Building, Adelaide : An 
architectural assessment'. 

No 20 ROWNEY, B. G. & DONOVAN, P. F. 
'Establishing conservation priorities for the huili 
environment; The example of Kapunda'. 



Ethnography 

PRETTY. G. L. 1969, The Maclcay Museum mummy 
from Torres Str.iils. a postscript ro iilliot Smith and 
the diffusion controversy. Man: the Journal of the 
Royal Anthropological Institute NS 4, 1: 24—43. 

PRETTY. G. L. 1969. 'Salvage Ethnography in New 
Guinea, the South Australian Museum Expedition to 
the Southern Highlands District, Papua, 1968-1969' 
Report to the Wenner-Gren Foundation for 
Anthropological Research, New York, iv + 85 pp.. 
illus. 



PRETTY, G L. 1970. Aboriginal Studies (Review of 
Man, Land and Myth in North Australia: the 
Gunwinggu people' by R. M. & C. H Berndt ; 
"Yiwara-Foragers of the Australian Desert' by 
Richard A Gould i Australian 8<mk Reviews % 12: 
332-334. 

PRETTY. G. L. 1974. Comment on Francis L. Utley, 
'The migration of folk tales: four channels to the 
Americas'. Current Anthropology IS, 1: 19-20. 

PRETTY, G, L 1980. The Aboriginal cultural 
landscape of the Lower Murray Valley. Pp. 43-52 in 
'Preserving Indigenous Cultures: i New Role for 
Museums'. Eds, R Edwards & J_ Stewart. UNESCO 
and Australia Council: Canberra. 

PRETTY, G L. (with A. CALDER). 1980. 
Mummification in Australian and Melanesia. 
Pp, 194-210 in 'Mummies, Disease and Ancient 
Cultures'. Eds. A, & E. Cockburn. Cambridge 
University Press; New York. 

PRETTY, G. L. and course participants. Visual Arts 
Discipline, Flinders University for the Gerard 
Aboriginal Communiry. 1983 'Artifacts of the 
Ngaiawang Tribe, Lower Murray Valley, South 
Australia, obtained by E. J. Eyre (1841-1844)'. 
(Document basis for the renewal of the Gerard 
Community artifact industry,) 

PRETTY 1 . G L. 1993 The Aborigines of the South East 
of South Australia and the Western District of 
Victoria before while contact. Community History 
(Adelaide) J, 4: 2-5. 



F.volutionary hiotogy 

PRETTY. G. L. (with P, SMITH & M. PROKOPEC). 
1988 Dentition of a prehistoric population from 
Roonka Flat, South Australia. Archaeology in 
Oceania 23; 31-36. 

rRKTTY. G. L. (with M PROKOPEC & P. SMITH). 
1994 Prehistoric South Australian Aboriginals: New 
evolutionary perspectives. Variability and Evolution 
(Adam Mickiewicz University. Poznan) 4. 21-56. 



Forensic anthropology 

PRETTY, G. L. |975. The recovery Of human remains 
lor forensic purposes, froceeilings Australian 
Forensic Science Society t, 3: 68-74 

PRETTY. G. JL (with D. h POUNDER & M. 
PROKOPEC), 1983. A probable case of euthanasia 
amongst prehistoric Aborigines at Roonka, South 
Australia. Forensic Science International 23: 99- 
1118 

PRETTY, G. L. (with M. PROKOPEC. D. SIMPSON 
& I. MORRIS). 1984. Craniosynostosis in a 
prehistoric Aboriginal skull: A case report Ossa. 
International Journal of Skeletal Research V-I1 
111-118. 



102 



H, TOLCHER. J. PRESCOTT & J MULVANEY 



Heritage preservation 

PRETTY, G L. 1967. 'Preliminary catalogue of 
archaeological monuments in the Territory Of Papua 
New Guinea'. Papua New Guinea Public Museum 
and Art Gallery: Port Moresby, jv + 95 pp., map 

PRETTY, G. L. 1968. Surveying archaeological 
monuments in Papua New Guinea. Journal of the 
Anthropological Society of South Australia 6. 2: 
5-8. 

PRETTY. G L 1970. Survey of prehistoric monuments 
in South Australia. Pp. 39-50 in 'Aboriginal 
antiquities in Australia: their nature and 
preservation'. Ed. F D. McCarthy. Australian 
Institute of Aboriginal Studies 22. 

PRETTY, G. L. 1976. The Ngaiawang Folk Province. 
Journal of the Anthropological Society of South 
Australia 14,9. 13-22. 

PRETTY. G. L. 1979. The Ngaiawang Folk Province. 
Pp. 28-45 tn 'The Tide of Ausualiun Scittcmeni 
Conservation of the Physical Evidence' Australia 
tCOMOS Proceedings, Beechwortb. 14-16 April 
1978. Auslralia/ICOMOS: Hawthorn, Vic. 



History 

PRETTY, G. L 1967 Edward Gibbon Wakefield. 
Pp. 559-562 in 'Australian Dictionary nf 
Biography" Ed. D. Pike. Melbourne. 

PRETTY, G. I. 1975. The historical collections 
in the South Australian Museum Pp. 173-176 
in 'Proceedings of the National Seminar on the 
Conscrvalion of Cultural Material. Penh, 
August 1973'. Eds C. Pearson & G. L. Pretty) 
Institute for the Conservation of Cultural 
Material: Perth. 

PRETTY, G. L. 1986 Australian history at Roonka. 

Journal of the Historical Societv of South Australia. 

14: 107-J22. 
PRETTY. G L. 1986 Foreword In People. Places and 

Buildings', J, Faull & G Young. South Australian 

Centre for Settlement Studies: Adelaide. 



Materials conservation 

PRETTY, G, L. (with C. PEARSON). (Eds). 3976. 
'Proceedings of the National Seminar on the 
Conservation of Cultural Material, Perth, August 
1973' Institute for the Conservation of Cultural 
Material: Penh. 297 pp., illus. 



Muscology 

PRETTY. G. L. 1967. The British Museum. Friends of 
the South Australian Museum Newsletter 4: 9-1 1 

PRETTY, G. L. 1968. Report on the Stale of the Papua 
New Guinea Museum: the Collections of 
Archaeology and Ethnology'. Trustees, Papua New 



Guinea Public Museum and Art Gallery: Pott 

Moresby, iv-f 95 pp., iiius. 

PRETTY. G. L. 1971 Oceanic museum resoutccs in 
Australian museums. Pp. 179-186 in 'Meeting on 
Studies of Oceanic Cultures' Australian National 
Advisory Committee for UNESCO: Canberra. 

PRETTY. G. L. 1976. The Ethnographic Collections in 
the South Australian Museum. Pp. 107-1 19 in 
'Proceedings, National Seminar on Conservation ut 
Cultural Material'. Eds. C Pearson & G L. Pretty 
Institute for the Conservation of Cultural Material: 
Perth. 



Palaeoecology 

PRETTY, G I. Uub W E BOYD). 1989. Some 
prospects tor archaeological palaeobotany in 
Australia; an example from South Austtalia. 
Australian Archaeology 28: 40 £2 

PRETTY. G L. (with W. E. BOYD), (in press). From 
[ha sensible to the spiritual - Cultural signals from 
palaeobotiiiin. evidence at Roonka, South Australia 
Archaeological Dialogue j 



Paleopathology 

PRETTY, G I.." (will. M. E. KRICUN) 1989 Health 

status of the prehistoiit Roonka population. World 

Archaeology 21, T, 198-224. 

PRETTY, C. L (with M. PROKOPEC) 1990 
Skeletal aging Rale of tooth attrition in hunter- 
gatherer populations, prehistoric Roonka, South 
Australia. Colloquiae Anthropalagkae 14, 2: 331- 

334. 

PRETTY. G. L. (with M. PROKOPEC). 1991. 
Observations on health, genetics and culture from 
analysis of prehistoric population data from Roonka, 
South Australia. Pp 151-158 in 'Human 
Palacopathology: Current Syntheses and Future 
Options', Eds. D. J. Ortncr & A. C. Aufdeiheide. 
Smithsonian Press: Washington, DC 

PRETTY. G. L. (with T. BROWN & M. E KRICUN). 
1992 Extensive compensatory remodeling of 
craniofacial structures; a case from prehistoric South 
Australia. Pp. 139-143 in 'Craniofacial Variation in 
Pacific Populations. Papers presented at the XVII 
Pacific Science Congress Symposium, Honolulu. 
Hawaii. May 30, 1991'. University of Adelaide: 
Adelaide. 

PROKOPEC, M and PRETTY. G. L. 1994. 
Praehistorische Bewohner der Roonka-Siedlung 
(Sudaustralien) Ethnographisch-Airharologische 
Zcitschrifl 35. 1: 136-148. 

SIMPSON, D., PROKOPEC. M.. MORRIS, t ... & 
PRETTY, G, L, 1984. Prehistoric craniosynostosis. 
A case report. Ret rods of the Adelaide Children's 
Hospitals, 2: 163-168. 



OBITUARY - GRAEME LLOYD PRETTY 



103 



Radiometric dating 

PRETTY, C. L. (with J. R. PRESCOTT, H. POLACH 
& B. W. SMITH). 1983. Some comparisons of C-14 
and thermoluminescent dates from Australia. 
(Proceedings of the 1981 Groningen C-14 
Conference). Council of Europe PACT Journal 8: 
205-211. 

PRETTY, G. L. 1988. Radiometric chronology and 
significance of the fossil hominid sequence from 
Roonka, South Australia. Pp. 32-52 in 'Early Man in 
the Southern Hemisphere'. Ed. J. R. Prescott. 
Supplement to Archaeometry: Australasian Studies. 
University of Adelaide Department of Physics and 
Mathematical Physics: Adelaide. 



Technology 

PRETTY, G. L. 1970. Review of 'Excavations of the 
prehistoric iron industry in West Borneo' by T. 
Harrisson & S. J. O'Connor. Man: The Journal of 

the Royal Anthropological Institute, London. NS. 5: 
710-711. 



Trace element chemistry 

PRETTY, G. L. 1991. (with F. D. PATE, J. T. 
HUTTON & R. A. GOULD). Alterations of in vivo 
elementary dietary signatures in archaeological bone. 
Evidence from the Roonka Flat Dune, South 
Australia. Archaeology in Oceania 26: 58-69. 



Helen Tolcher, c/- South Australian Museum, North Terrace, Adelaide, South Australia 5000; Professor Emeritus 
John Prescott, Physics Department, Adelaide University, South Australia 5005; and Professor Emeritus John 
Mulvaney, Centre for Archaeological Research. Australian National University, ACT 0200. Records of the South 
Australian Museum 35(1): 97-103. 



imTni^J 



SOUTH 

AUSTRALIAN 

MUSEUM 

VOLUME 35 PART L. 
MAY 2002 
ISSN 0376-2750 

CONTENTS: 



ARTICLES 

1 1. BEVER1DGE 

New species and new records of Cloacina von Linstow, 1898 (Nematoda: Strongyloidea) 
parasitic in macropodid marsupials from Papua New Guinea. 



33 B. P. KEAR 

Darwin Formation (Early Cretaceous 
Australian Museum. 



reptile remains in the Sout 



49 H. W. PALM & I. BEVERIDGE 

Tentaculariid cestodes of the order Trypa 
Australian region. 



(Plutyhelminthes) from the 



79 E. G. MATTHEWS & T. A. WEIR 

T "'o new species of the genus Lepanus Balthasar from South Australia (Coleoptera: 
Scarabaeidae). 

85 S. BARKER 

A checklist of Cisseis (sensu stricto) Gory & Laportc, 1839 (Coleoptera: Buprcstidae: 



91 W. ZE1DLER 

The Flora and Fauna of South Australia Handbooks Committee 10 March 1921 - 
30 October 200 1. 



97 H. TOLCHER, J. PRESCOTT & J. MULVANEY 
Obituary - Graeme Lloyd Pretty 



Published by the South Australian Museum. 
North Terrace. Adelaide. South Australia 5000. 



THE LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 

(COLEOPTRA : INSECTA) 

C. H. S. Watts 



Summary 

The larvae of the aquatic Hydrophilid genera Allocotocerus Kraatz, Hybrogralius Orchymont and 
Regimbartia Zaitzev are described and figured for the first time.The larvae of the following 
Australian species are also described and figured, most for the first time: Allocotocerus punctatus 
(Blackburn); Amphiops aueenslandicus Balfour-Browne; Berosus australiae Mulsant; Enochrus 
eyrensis (Blackburn), E. mastersi (W. MacLeay); Helochares tristis (W. MacLeay), H. clypeatus 
(Blackburn), H. tenuistriatus Regimbart, H. foveicollis (Montrouzier); Hydrobiomorpha sp. ; 
Hydrophilus bilineatus (MacLeay); Hybrogralius hartmeyeri (Regimbart); Laccobius decipiens 
Gentili; Limnoxenus zealandicus (Broun); Regimbartia attenuata (Fabricius); and Sternolophus 
margincollis (Hope). 



THE LARVAE OF SOME AUSTRALIAN AQUATIC 
ITYDROPHILIDAE (COLEOPTERA: INSECTA) 



CHS WATTS 



WATTS, CHS 2002. The larvae of some Australian aquatic Hydrophilidae (Coleoptcra: 
Insccta), Records ql the South Australian Museum 35(2): 105-138. 

The larvae of the aquatic Hydrophilid genera Allocotocerus Kraatz, Hybugralius Orehymonl 
and Regtmbarlia Zaitzev are described and figured for the first time. The larvae of the 
following Australian species arc also described and figured, most for the firs! time: 
Allocotocerus punctatus (Blackburn); Amphiops aueenslandicus Balfour-Browne; Berosus 
australiae Mulsant; Enochrus eyrensis (Blackburn), E. maslersi (W. MacLcay)', Helochares 
tristis (W. MacLeay), H. luridus (W. MacLcay), //. clypeatus (Blackburn), H. tenuislrialus 
Regimbart, H. foveicoths (Montrouzicr); Hydrobiomorpha sp.; Hydrophilus bilineams 
(MacLeay), Hybogralius hartmeyeri (Regimbart); Laccobius decipiens Centili; Limnoxenus 
zealandicus (Broun); Regtmbarlia attenuaia (Fabricius); and Strrnuliiplms murginieollis 
(Hope). 

The larvae were identified by rearing larvae collected in the field, or occasionally by 
association and elimination, or by using biochemical methods. 

The pupae of Amphiops qucenslandicus and Hybogralius hartmeyeri are described and 
figured. Unlike most other Hydrophilids the larvae of Amphiops arc thought to pupate exposed 
on the stems of emergent vegetation. 

A key is provided to the larvae of Australian genera of aquatic Hydrophilids other than 
Notahxdrus Balfour-Browne, Paranacaena Blackburn. Phelea Hanson and Agraphydrus 
Regimbart, whose larvae remain unknown. 

CHS Watts, South Australian Museum, North Terrace. Adelaide. South Australia 5000 
Manuscript received 23 April 2001. 



The Hydrophilidae (serisu Hansen 1991) 
comprise a major part of lhe Australian aquatic 
beetle fauna, occurring in all types of fresh water 
and most commonly among vegetation at the edge 
of standing water. Adults and larvae occur 
logeiher. The taxonomy of the adults is now 
relatively well known thanks to the work of 
Hansen (1991) at the generic level and above; and 
that of Gentili (1980. 1992. 1993, 2000) and 
Walts (1987, 1988, 1989, 1990, 1995. 1998a, 
1998b) on the aquatic fauna at the species level. 
No recent work has been done on the species level 
taxonomy of the extensive portion of the family in 
which both aduhs and larvae are terrestrial. 

In contrast to the situation with adults, lhe only 
work specifically on Australian larvae is that of 
Anderson (1976) who described the larvae of 
Helochares tristis (W MacLeay). Enochrus 
maculkeps (W MacLeay) and Chasmngenus 
nitescens (Fauvel) together with details of iheit 
life histories. 

One feature ul the Australian aquatic 
Hydrophilid fauna is its low endemicity at the 
generic level. Consequently, descriptions of a 



number of Australian genera are available in 
scattered Northern Hemisphere publications. The 
work on the New World genera was summarised 
and added to by Archangelsky (1997). This work 
included a number of genera which also occur in 
Australia. 

For a number of years I have been rearing field- 
collected larvae of Australian aquatic 
Hydrophilids with the aim of discovering and 
describing the larvae of all genera and, for those 
already known from non-Australian species, to 
extend the descriptions to incorporate Australian 
species. The basic motivation was to produce a 
key to enable the larvae of all Australian aquatic 
Hydrophilidae to be identified at least to genus. 
As a result I have reared 12 species in 9 genera. 
Two additional genera, Allocotocerus and 
Regimbartia, were identified by associating adults 
and larvae by the biochemical technique of 
altoen/yme electrophoresis. Others were identified 
using the descriptions in Archangelsky (1997), eg 
Hydrobiomorpha; or by association and 
elimination, eg Enochrus maslersi. 

I have not attempted any phylogenetic analysis 



106 



CHS WATTS 



since, with such a high proportion of genera also 
occurring widely outside Australia, the only 
sensible approach would have been to include an 
examination of all known genera, which was 
beyond the scope of this study. Some comments 
on relationships are given under most generic 
descriptions. 

In preparing this paper I have drawn heavily on 
die work of Archangelsky (1997) on the New 
World fauna. The descriptions follow his format 
and a number of the figures used to illustrate the 
key come from hi>> importanl publication. 

Although their larvae are known from other 
places, 1 have not seen Australian examples of the 
following genera Chaetarthria Stephens, 
Chasmogenus Sharp, Crenitis Bedel. Coehstoma 
Brulle or Paracymus Thomson, and since 1 have 
nothing to add have not included descriptions. 
Detailed descriptions can be found in 
Archangelsky (1997). In addition, the larvae of 
Paranocaena Blackburn, Nolahydrus Balfour- 
Browne, P he lea Hanson and Agraphydrus 
Rcgimbart are unknown. 



Materials and Methods 

Larvae were collected in the field and reared in 
small aquaria formed hy placing the base of a 
small pclrie-dish inside a larger petrie-dish and 
filling the space between them with damp sand 
(Walts 1963). Mosquito and chironomid larvae 
were the main food items provided. These were 
accepted by all larvae other than those of 
Hydraphtlus (see under HydruphUus). All species 
successfully reared pupated in cells constructed in 
the wet sand with the exception of those of 
Amphiops which pupated on the surface or 
attached to the sides of the container. 

Larvae were preserved in 75% ethanol and 
examined under a slereomicroscope. Permanent 
microscope slides were prepared for detailed 
examination. Drawings were made with the aid 
of a camera lucida Habitus drawings were only 
prepared when no previous illustration was 
available or when Australian species differed 
from those previously illustrated Unless 
otherwise mentioned the descriptions arc based 
on third instar larvae. Earlier instars usually 
differ somewhat, most noticeably in the stouter 
cephalic appendages and in fewer spines on the 
menlum. The key is constructed to accommodate 
all instars. Measurements of the head capsule 
were made under a stereoscopic microscope wilh 
a graduated eyepiece. Measurements of total 



length were made with dial callipers on hoi li 
mounted and alcohol preserved material, but in 
this case the measurements can only be 
indicative as factors such as larval age and 
preservation effects can alter the length 
considerably. Measurements of the head capsule, 
particularly the width, are a more reliable 
indicator of size Unless slated otherwise, the 
specimens were collected by myself and arc in 
the collection of the South Australian Museum. 



System attcs 

KEY TO THE GENERA OF AUSTRALIAN 
AQUATIC HYDR0PHIL1DAE BASED ON- 
LARVAL CHARACTERS 

1) Not including the genera Notohydru\. 
Puranaiaetui, Phelea and Agraphydrus, 
whose larvae are not known. 

2) tneluding Spercheus which is in the family 
Spercheidae but which has larvae that can be 
confused with Hydrophilidae. 

1 — Tip of mandible bifid (Fig, 2g) 

i... Spercheidae (Spercheus) 

— Tip of mandible not bifid 

Hydrophilidae... 2 

2 — Hypopharyngcal lobe well developed, like 

a pubescent tongue, originating at the base 
of the labium on the left side (Figs 2a, 2b) 

Subfamily Sphaeridiinac* 

Coelosumia Brulle 

— Hypopharyngcal lobe reduced or ahseni 
Subfamily Hydrophilinae. 3 

3 — Abdominal segments with long 

filamentous gills (Fig, 5a i, or with multiple 
setose lateral projections on abdominal 

and thoracic segments (Fig. 15a) 

Tribe Bcrosini...4 

— Lacking long filamentous abdominal gills 
or complex lateral projections, at most 
with simple shorl fleshy projections (Figs 
4a, 14a, 16a) 6 

4 — Gills long, without setae (Fig. 5a); 

labroclypcus asymmetric (Fig. 5b). 
mandibles asymmetric (Fig. 5f); 
prementum small and squat, without ligula 
(Fig 5c), basal segmentol anlenna without 

laicral projection (Fig. 5e) r 

Berosus Leach 

Abdominal and thoracic segments each 
wuh iwoormore setose, lateral projections 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAF 



107 



7 — 



8- — 



(Fig 15a); mandibles nearly symmetrical 
(Fig. ! 5h); prememum long and thin, with 
ligula(Fig 15d); basal segment of antenna 
wiih lateral projection (Fig. I5g) 5 

Body spines predominantly blunt, weakly 
bifid at lips (Fig 3b); lateral projection on 
basal segment of antenna nearapex, sharp. 
spine-like (Fig. 3f);apex of second segment 
of antenna with lateral extension (Fig. 3d); 
maxillary stipe without spines (Fig. 3e); 
ligula relatively short, prememum stouter 

(Fig. 3c) AllocuWcerus Kraatz 

Body spines predominantly pointed (Fig. 
15c); lateral projection on basal segment 
of antenna thumb-like., further from apex 
(Fig. 15g), second segment of antenna 
without lateral extension (Fig. 15e); 
maxillary stipe with spines on inside (Fig. 
I5f); ligula relatively long, prementum 

more elongate (Fig. 15d) 

...Regimbartin ZaiUev 

Apical segment of antenna about same 
length as penultimate (Figs 1 2d, I6e); 
femora with fringeof swimming-hairs (Fig- 
2h); withprosty lesflwo retractable, fleshy 
appendages on last abdominal segment) 

Subtribe Hydrophilina...7 

Apical segment of antenna much shorter 
than penultimate (Figs 9e, 14e); femora 
lacking, or virtually lacking, swimming- 
hairs; without prostyles 9 

Labroclypeus without teeth (Fig. 12a); 
mandibles asymmetrical, left one very 
robust, right one more slender (Fig. 12e): 
ligula shorter than lirst segment of labial 
palpus (Fig. 1 2b); up lo 40 mm long .... 

Hytltvphilus Muller 

Labroclypeus with weak to moderate teeth 
(old specimens may lacktccthdue to wear) 
(Figs Me, 1 6f); mandibles symmetrical or 
nearly so; ligula longer than first segment 
of labial palpus (Figs lib, 1 6c); up to 
20 mm long 8 

Prementum longer than mentum (Fig. 1 6c), 
first segment of antennae with few spines 
(Fig, 1 6f); mandibles each with two large 
distal teeth and one small proximal denticle 

(Fig 16f) ..Sternnlophus Solier 

Prementum slightly shorter than mentum 
(Fig. 1 lb); first segment of antenna with 
numerous spines on inside (Fig. lie); 
mandibles each with one large distal tooth 
which is pick shaped at the tip and one or 



10 



II — 



\2 



13 



14 



two small proximal denticles (Fig. I le) 
Hydrabiomarpha Blackburn 

Left expansion of epistoma much more 
prominent than the right and widi a row of 
stout setae on front edge (Fig. 1 3b); left 
mandible with group of stout setae at base 
of middle tooth which is lacking on the 

right mandible (Fig 130 10 

Lateral expansions of epistoma similar, 
without a row of stout setae on front edge 
(Fig. 14b); mandibles without such a group 
of setae II 

Frontal sulci parallel and not uniting to 
form a coronal sulcus (Fig. J3a); nasale 
with three teeth (Fig. 13b); prementum 
wider than long (Fig 13c); ligula absent or 

virtually so (Fig. 13c) 

Laccobiits Erichson 

Frontal sulci meet just before occipn.il 
foramen to form a coronal sulcus (Fig. 
10a); nasale with five teeth (Fig. 10b); 
prementum longer than wide (Fig. 10c), 
ligula present though small (Fig. 10c) .. 
,„,.„ Hybogmlius Orchymont 

Legs very short, without claws, prementum 
with a large round ligula (Fig. 2e); length 

up to 6 mm Clmetarthria Stephens 

Legs usually longer, with claws. 
Prementum wiih or without ligula, but if 
present never large and round , length up to 
15 mm I? 

Mentum large, widerthan long, prementum 

small, lacking ligula (Fig. 4b) 

Amphiops F.richson 

Mentum square or longer than wide, 
prementum well developed, ligulaprescnt 
(Figs 6b, 8b, 14c) 13 

All abdominal segments with dorsal 
sclerites, although often small; without 
coronal sulcus (Fig. 2c); length upto6 mm; 
antennal appendage as iong as apical 
segment; mandibles symmetrical with three 

teeth Tribe Anacae.nini.,,,14 

Abdominal segments, other than I and 8, 
lacking dorsal sclerites; with short coronal 
sulcus (Fig, la); antennal appendage hall 
length of apical segment (Fig. 8d); 
mandibles often asymmetrical and with 
fewer than three teeth, length up to 15 mm 

..... 15 

With lateral abdominal flaps (Fig. 2c); 
without cervical sclerites Crenitis Bedel 



108 



CHS WATTS 




-9 
-10 

-11 

-12 

-13 
-14 




FIGURE 1. a, dorsal side of head capsule of Hybogralius hartmeyeri. 1, maxillary palpus; 2, stipes of maxillary 
palpus; 3, mandible; 4, mandibular teeth; 5, labium; 6, antenna; 7, right lobe of epistome; 8, teeth of nasale; 9, 
nasale (7 & 9 make up the labroclypeus); 10, stemmata; 11, frontal sulci; 12, coronal sulcus; 13, cervical sclerites; 
14, occipital foramen; b, details of dorsal side of labium of Hybogralius hartmeyeri. 1, labial palpus; 2, ligula; 3, 
prementum; 4, mentum. 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHIL1DAE 



109 






FIGURE. 2. a, ventral view of labium and hypopharyngeal lobe (arrowed) of Sphaeridium .uarabaeoides 
(Linnaeus) (subfamily Sphacridiinae); b, dorsal view of labium and hypopharyngeal lobe (arrowed) of 
Daaylosternum sp. (subfamily Sphacridiinae); c, habitus of Crenilis morata (Horn); d, ditto, dorsal view of 
labium; e, ditto of Chaetarthria sp., f, ditto, Paracymux suhcupreus (Say); g, mandible of Spercheux platycephalux 
MacLeay; h, hindleg of Hydrobiomorpha sp., Northern Territory; i. labroclypeus and mandible of Chasmogenus 
nitescens (Fauvel). a-f, North American laxa from Archangelsky 1997; i. from Anderson 1976. 



110 



CHS WATTS 



Lfi 



17 — 



— Lacking abdominal Haps; with small 
cervical sclerites . Paracymus Thomson 

l*i — Apical segment of antenna about halt 
length of penultimate (Fig. 14e); mandibles 
symmetrical with three teeth, without 

serrations (Fig. I4f) 

Limnnxeims Motschulsky 

— Apical segment of antenna about a quarter 
the length of the penultimate segment 
(Fig. 8d); mandibles weakly to strongly 
asymmetric, with one or two teeth, often 

with serrations (Figs 6e, 8ej 

Subtrihe Acidocerenia....l6 

Nasale moderately developed, with right 

side more prominent (Fig. 6a> 17 

Nasale poorly developed, all areas equally 
prominent (Fig. 9b) 19 

Mandibles asymmetric, the right one 
with one tooth, left with two (Fig. 6e) 

Enochru:. 

Thomson (other than E. mastersi) 

• — Mandibles almost symmetrical, both with 

two teeth (Fig. 8e) 18 

IS — Inside of maxillary stipe with group of 
spines near base (Fig. 8c); ligula longer 
than first segment of labial palpus (Fig. 
8b); prementum approximately the same 
length as mentum (Fig Kb). ..Helochares 
Mulsanl (other than H. foveicalli.t) 

— Inside of maxillary stipe lacking such 
spines (Fig. 7c); ligula shorter than first 
segment of labial palpus (Fig. 7b); 
prementum approximately twice length of 

mentum (Fig. 7b) 

E, mastersi (W, MacLeay) 

19 — First segment of antenna with bulge on 
inside near apex (Fig 9e); mandibles each 

with IwotccthfFig. 80 

.H. foveicollis (Monirourier) 

— First segment of antenna without bulge; 
right mandible with one tooth, left one 

with two (Fig. 2i) 

Chasmogentts Sharp 

"* Members of the subfamily Sphaeridiinac are 
predominantly terrestrial, occurring in moist 
situations such as dung and rotting vegetation. 
Members of only one genus, Coelostoma, are 
commonly found in aquatic situations in 
Australia. 



Descriptions 

The following descriptions are arranged in 
alphabetic order of the genera. 



Allocatocerus Kraatz 

Allocatocerus punctatus (Blackburn) Fig. 3 

Size of third instar. Length 8.5-12.0 mm, head 
capsule, 0.70-1.00 mm long, 0.85-1.0 mm wide. 

Head capsule. Subquadrangular, bulging 
outwards at anterolateral angles. Labroclypeus 
symmetrical; nasale very short, with numerous 
very short teeth on anterior border of epislome; 
lateral lobes of episiotne rounded, very short. 
Frontal sulci straight, reaching from antennal 
bases to rear of head. Coronal sulcus absent Gular 
sclerile absent. Cervical sclerites small, 
suhrectangular. very weakly sdcrutiscd. 

Antennae. Three-segmented. First segment 
slender, longer than other two combined, with a 
spine-like inner process close to apex; third 
antennal segment short, connected to edge of the 
second segment before apex. Sensory appendage 
on second segment, slim, shorter than third 
segment. 

Mandibles. Symmetrical, long and slender, with 
two inner iceth on basal half, distal tooth large, 
basal one small. 

Maxillae. Five-segmented. Stipes slender, much 
longer than remaining segments combined, with 
two or three fine setae on inner margin. Palpus 
four- segmented, first and third segments subequal 
in length, first segment bearing a short inner 
process; second segment very short; fourth 
segment shorter than third. 

Labium. Mentum large, subglobular, dorsal 
surface wilh some cuticular spines towards sides, 
anterolateral angles rounded wilh a few spines. 
Prementum much longer and narrower than 
mentum, suhrectangular, longer rhan wide. Palpus 
I wi -i- segmented, basal segment short. Ligula about 
two limes as long as basal segment of palpus, tip 
broadened 

Thorax and legs. Prolhorax wilh broad dorsal 
sclerite, with saginal line, surface covered by 
minute spines, fringe of long, slender setae around 
the margins and three short setae-bearing 
projections at sides; ventral surface with large, 
subreclangular sclerite, without sagittal line. 
Mcsothorax wilh two dorsal pairs of subtriangular 
sclerites Meiathorax wilh one dorsal pair of 
uregular shaped sclerites. Both mesothorax and 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 



111 




FIGURE. 3. Allocotocerus punctatus. a, labroclypeus; b, detail of abdominal Stella; c, labium; d, tip of antenna; e, 
maxilla; f, antenna; g, mandibles. 



112 



CHS WATTS 



metathorax with three lateral pairs of setiferous 
projections and a number of short setiferous 
papillae on dorsal surface. Legs five-segmented, 
long, visible from above. 

Abdomen. Segments 1 to 7 with four pairs of 
setiferous projections similar to those on thorax, 
three on the side and a smaller dorsal one 
towards the midline. In addition, the abdomen is 
covered with minute, simple spines and the 
dorsal surface with small projections, each with 
six to eight, relatively long setae, truncated or 
weakly bifid at tips (in most specimens these 
structures are obscured by an accumulation of 
sand grains and detritus). Segment 8 without 
setiferous projections; with dorsal semispherical 
sclerite covering the spiracular atrium; with a 
pair of small finger-like 'procerci'. Segment 9 
trilobed, with a pair of small, unsegmented 
urogomphi. 

Spiracles. Mesothorax and abdominal segment 
2 with pair of small non-functional spiracles on 
short papillae. (I have been unable to detect 
spiracles on other abdominal segments.) Spiracles 
on segment 8 annular, large and functional, within 
the spiracular atrium. 

Interspecific variation 

There is little variation among the few 
specimens seen — including one from the 
Northern Territory which is almost certainly A. 
tibialis (Balfour-Browne). 

Identification 

By association of adult and larva by isoenzyme 
electrophoresis. 

Remarks 

The larvae closely resemble those of the New 
World genus Derallus (Archangelsky 1997; 
Spangler 1966) in the setiferous body structures, 
form of the labroclypeus, mandibles, labium, 
maxillary palpi and antennae. The differences arc 
small: blunt rather than sharp-pointed body spines, 
shape of the distal part of the antennae, number of 
spines on the maxillary palpi and details of the 
labium. The larva of Allocotocerus is also closely 
similar to those of Regimbartia, differing from 
this genus in the details of the same suite of 
characters. On balance it more closely resembles 
Derallus. Although placed by Hansen (1991) with 
Berosus in the tribe Berosini, on larval characters 
the three genera form a very distinctive and 
cohesive group well separated from Berosus, a 
conclusion reached by others (Bertrand 1972; 
Oliva 1992; Spangler 1966). 



Specimens examined 

Queensland: 2 km S Mt Molloy, 30/3/96. 
Allocotocerus sp. Northern Territory: Manton 
Dam, 23/3/98. 



Amphiops Erichson 

Amphiops queenslandicus Balfour-Browne. Figs 
4, 17a 

Size of third instar. Length 6.5-9.0 mm; head 
capsule 0.65-0.80 mm long, 0.78-0.82 mm wide. 

Head capsule. Subquadrate. Labroclypeus 
symmetrical; nasale very short, with five small 
teeth; lateral lobes of epistome rounded, 
projecting further than nasale. Frontal sulci 
inversely bell-shaped, meeting before reaching 
occipital foramen. Coronal sulci very short. Gular 
sclerite absent. Cervical sclerites small, 
rectangular. 

Antennae. Three-segmented. First segment as 
long as second; third half the length of second. 
Sensory appendage on second segment short, a 
quarter the length of the third segment. 

Mandibles. Symmetrical with three inner teeth, 
the two distal teeth large and with slightly 
crotchet-shaped tips; third tooth much smaller. 

Maxillae. Five-segmented. Stipes narrow, 
longer than the remaining segments combined, 
with a row of five to six small setae on inner 
margin. Palpus four-segmented; first segment 
elongate, rectangular, with inner process as long 
as very short second segment; third segment 
longer and narrower than first; fourth segment a 
little longer than second. 

Labium. Mentum large, subrectangular, anterior 
edge with five to six large spines/protuberances, 
and numerous cuticular spines at sides towards 
base. Prementum ovoid, much smaller than 
submentum. Palpus short, two-segmented, basal 
very short, distal segment two to three times 
longer than basal one. Ligula absent. 

Thorax and legs. Pro-, meso- and metasterna, 
almost completely covered by dorsal sclerites, 
each with a sagittal line. Ventral surface of 
prothorax with a subrectangular sclerite, 
subdivided by a sagittal line. Sclerites with very 
small spines arranged in short lines towards rear 
(only visible under high magnification). Bases of 
setae on sclerites pigmented. Legs five-segmented, 
moderately developed, visible in dorsal view. 
Dorsal surface with sparse covering of small dark 
patches. 

Abdomen. First segment with dorsal pair of 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPH1LIDAE 



113 









a d e f 

FIGURE 4. Amphiops qutinslanduus. a, habitus; h, labroclypeus; c, labium; d, maxilla; e, anicnna; f. mandible 



moderate sized subtriangular sclerites, segments 2 
to 5 with smaller dorsal sclerites, sclerites with 
scattered small darker patches. Segments 1 to 7 
similar in size and shape, subdivided by a 
transverse fold; pleural areas with four pairs of 
moderate bulges/lobes each with several long 
setae at apex, ventral one largest; with a pair of 
small dorsal papillae, each with long terminal 
setae, abdominal surface strongly folded. Segment 
8 with pair of small 'procerci', with a large dorsal, 
suboval sclerite; segment 9 Irilobed. with a pair of 
small one-segmented urogomphi. Integument with 
only a few small spines. 

Spiracles. Eight abdominal pairs; pairs 1 to 7 non- 
functional, on tips of small papillae and each with a 



long seta immediately inwards on a small cylinder- 
shaped base. Spiracles on segment 8 annular, large 
and functional, within the spiracular atrium. 

Pupa. (Fig. 17a) 

Colour. Light to dark brown. 

Head. With one pair of supraorbital styli; styli 
without apical setae. Appendages adpressed 
tightly to body, weakly differentiated externally. 

Thorax. Pronotum with 16 styli, all on margins 
of pronotum. Mesothorax lacking obvious styli; 
metanotum with two pairs of styli, lateral ones 
near the base of wingpads very small, central pair 
prominent. Styli without setae. Wing cases 
grooved/ridged. Legs held tightly against body 
almost totally covered by ptcrolhccac. 



114 



CHS WATTS 



Abdomen. Segments t to 7 with a transverse 
row of four to five very small tergal styli on a 
sharp ridge that is stronger laterally. Segments I 
to 7 with well-developed rigid (lap-like lateral 
extensions, those on segments 1 to 3 upright and 
slightly bent over inwards, those on segments 4 to 
7 horizontal and each with a small stylus on 
posterior edge, eaeh stylus with a small seta. 
Anterior edge of first tergite with very prominent, 
thin, rigid ridge. Segment 8 without obvious styli. 
segment 9 bearing a pair of medium-sized eeiei. 

Interspecific variation 

There is little variation among the specimens of 
Amphiops available, although it is almost certain 
that several species are included. There are some 
differences, namely the extent of the dorsal small 
dark patches/spots varies from a few tc moderate 
numbers; the area between ihe abdominal sclcritcs 
and spiracles sometimes is a bit darker than rest, 
and there is some variation «n the width of the 

nas&le. 

Identification 

By rearing larvae collected tn the field, 

Remarks 

Bertrand ( 1 972) keyed out Amphiops larvae and 
briefly illustrated a presumed larva. Australian 
species appear to have less well-developed lateral 
abdominal protuberances than the specimen from 
the Indonesian Archipelago (Insulinde) illustrated 
by Bertrand (1972) Berge Henegouwen (1982) 
described an African species as. having short 
conical outgrowths. 

The noticeably lumpy sides to the abdomen, the 
very small prcmenturri and labial palpi, and the 
I.Kk of a ligula most readily identity specimens of 
Antphiaps. 

The form of the pupae is unusual, in particular 
the virtual absence of setae, the compact ventral 
surface and hard shield-like dorsal surface. 

In the laboratory A. queenslanxticus (and also A 
duplopunctatus, of which a single larva was 
reared through to adult but the larval exuvium was 
lost) pupated above ground in the aquaria. This, 
together with the unusual shape and dark colour 
Or the pupae, suggest that in the wild they pupate 
attached to the stems of emergent vegetation. Just 
how they physically attach themselves remains to 
be discovered. 

Specimens examined 
Queensland I km W Mingela, 4/5/98. 
Amphiops sp. Northern Territory Jabiru, 



21/3/98; 11 km E Jabiru, 19/3/98; Manton Dam, 
23/3/98; Omiision Gorge, 27/1/99. Queensland: 
10 km S Cardwcll, 7/2/97; 40 km S Cardwell. 
7/2/97; 5 km NE Mt Molloy, 30/3/96, 2 km S Mt 
Molloy 30/3/96; Nardello's Lagoon near Mareeba. 
29/3/96; same locality, 6/2/97; 10 km W Pctric, 
23/1 1/95; 1 2 km NW Townsville. 8/2/97; 25 km 
S Townsville. 25/3/96. 



Berosus Leach 

Berosus australlae Mulsant. Fig 5 

Size of third itistar. Length 6.5-1 .05 mm 
(exclusive of gills); head capsule 0.75-0.95 mm 
long. 0.85-1 02 mm wide. 

Head capsule.. Subquadrate. Labroclypcus 
strongly asymmetrical; nasale narrow, strongly 
protruding, left of centre, with five to seven short 
teeth; lateral lobes of epistomc asymmetrical, right 
one weakly projecting, left one strongly projected 
•interiorly, covering basal third of mandible, with 
several short, strong, curved spines. Frontal sulci 
lyrifortn, not coming together, absent on third 
insiar larvae. Coronal sulcus absent. Gular 
sclerites absent, a pair of small, weakly 
sclerotised, cervical sclerites. With inverted V- 
shaped darker markings on dorsal surface. 

Antennae. Three-segmented, relatively close 
together, arising just inwards of mandible bases 
First segment a little longer than other two 
combined, with a stout subapical seta on inner 
margin. Sensory appendage on second segment, 
about half length of short third segment 

Mandibles. Slender, strongly asymmetrical; 
right mandible with large distal tooth and a very 
small central one; left mandible with group of 
seven small teeth or strong spines, some of which 
are bifid at tips. 

Maxillae, Five-segrnented. Stipes much longer 
than remaining segments combined, with four or 
five setae on inner margin. Palpus four- 
segmented; first segment short, subquadrate, with 
an inner process, second segment very short; third 
segment the longest; fourth a little longer than 
first. 

Labium. Mentum short, poorly sclerotised. 
Prernetitum small, subquadrate. Palpus long, two- 
segmented, basal segment shorter than distal. 
Ligula absent or reduced to small bulge. 

Thorax and legs. Prothorax with large dorsal 
sekritc, with sagittal line; ventral surface with a 
large subrceuingular sclente, hind edge strongly 
sinuate, with sagittal line. Mesothorax with two 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHIL1DAE 



115 




a d e f 

FIGURE 5. Berosus austmliae. a, habitus; b, labroelypeus; c. labium; d, maxilla; e, antenna: f, mandibles. 



dorsal pairs of sclcritcs, inner pair small, outer 
pair much larger, subtriangular. Metathorax 
lacking sclerites. Mesothorax and metathorax with 
a pair of small lateral tubercles bearing a long 
apical seta. Legs five-segmented, long, visible in 
dorsal view. 

Abdomen. Abdominal segments 1 to 7 
subdivided by a transverse fold; each with a pair 
of long, lateral, tracheal gills, and two pairs of 
small cylindrical structures on dorsal surface in a 
line inward from spiracles. Segment 8 small, with 
a dorsal sclerite; spiracular atrium absent. 
Urogomphi reduced. Integument densely covered 



with small setae. Bases of gills with covering of 
small setae similar to those on adjacent abdominal 
segments. Each gill with a relatively long seta, 
arising from a small cylindrical ba.se, towards tip. 
Dorsal surface often with darker patterning. 

Spiracles. Nine pairs of non-functional 
spiracles, one mesothoracic and eight abdominal. 

Intraspecifw variation 

There is some variation in the specimens I have 
identified as this species in the number and shape 
of the nasale teeth and in the presence/absence/ 
strength of a dorsal colour pattern. 



116 



CHS WATTS 



Interspecific sanation 

The description of.fi. austratiae fits mosi of 
the specimens of Berosus that I have seen, 
many of which are unlikely lo be B. austratiae. 
1 suspect that the description would fit the 
Larvae of most species in the B austratiae, B 
majusculus Blackburn, B. veronicae Watts 
group of Australian Berosus. There is some 
variation in colour and in the strength of setae 
on the integument (weak in B austratiae). 
Another group of specimens show clear 
differences from B austratiae: the nasals is 
central rather than to the left; the integument 
is covered with small dark-pigmented patches 
in various patterns; the abdominal segments 
have variably sized, but often marked, colour 
pallern; there are finger-like projections on the 
sides of the prothorax and a differently shaped 
eighth abdominal segment. Those larvae are 
associated with species such as B. 
queenstandicus Blackburn and B involutus 
(W. MaeLeay). 

Identification 
By association and elimination. 

Remarks 

The long filamentous lateral gills and the 
absence of functional spiracles readily identify 
larvae of Berosus. The asymmetry of the 
mandibles and labrodypcus, stoat maxillae 
and lack of a ligula are also distinctive, but are 
characters shared variously by a number of 
other genera Wilhin Australian species there 
is no sign Of the variation in the number of 
pairs of lateral gills seen in African (Bcrlrand 
1972) and New World (Archangelsky 1*397) 
species. 

Specimens examined 

Queensland: Bohle River. 10 km N 
Townsville, 23/3/96 

Berosus sp. New South Wales: S km N 
Bombala, 28/11/98. ditto. 3/11/97; 
Collector. 20/1/97; 5 km NE Dartmoor, 
11/10/96. Northern Territory: Mt 
Borradaile Station. 26/5/99; Nawurlandja, 
Kakadu National Park, 29/1/99. 
Queensland: Burdekin River, 2/1 1/95; 10 K 
S Cardwell, 7/2/97. Jourama Falls. 31/10/95; 
5 km NE Mt Molloy, 30/3/96. South 
Australia: 10 km N Coonawarra; 19/10/9') 
Victoria: Simpson Creek, 12 km SW Orbost, 
16/1/97. Western Australia: Gin Gin, 
15/10/96; 4 km S New Norcja. 15/10/96. 



Enochrus Thomson 

Enochrus eyrensis (Blackburn), Fig. 6 

Size of third instar Length 9.1 mm; head 
capsule 0.70-0 75 mm long; .0.75-0.80 mm wide 

Head capsule Subquadratc. Lahroclypeus 
asymmetrical; nasale obliquely truncate, left side 
shorter than right, with seven to nine short teeth, 
first two on right side and last one on left side 
largest; left lobe of epistome rounded, shorter than 
nasale; right lobe triangular, sharply pointed, 
about same length as nasale Frontal sulci as an 
inverted bell, fusing just before reaching occipital 
foramen. Coronal sulcus very short Gular scleriie 
absent. Cervical scleritcs moderate, 
subrcctangular. 

Antennae. Three-segmented. In third instar firsl 
segment wider than second segment and about 
same length; second segment with small setae - 
beanng appendage on inside near middle: third 
segment about a third the length of the second. 
Sensory appendage on second segment much 
shorter than third segment. 

Mandibles. Strongly asymmetrical; right 
mandible with two strong inner teeth on basal 
half; left mandible with one strong inner tooth. 
Inner margin or right distal tooth and distal parts 
of both mandibles serrated, 

Matiltae. Five-segmented. Stipes wide, much 
longer than remaining segments combined, with 
four strong setae on inner margin Palpus four- 
segmented; first segment subrectangular, a little 
wider that long, with an inner process a little 
shorter than second palpal segment: second 
segment short; third and fourth segments about 
twice as long as second, subequal in length. 

Labium. Mentum subtrapezoidal, dorsal surlac: 
with two short stout spines in middle towards 
fronl. three strong spines laterally, anteriolaieral 
angles each with a strong spine Prcmcntum squat, 
a little wider at base, shorter than menlum. Palpus 
two-segmented, first segment short, second 
segment approximately twice as long as first. 
Liguta slender, longer than first palpal segment, 
with partial rinj> of small spines around bases, of 
palpal segments. 

Thorax and legs. Prothorax completely covered 
by a dorsal sclerite, with sagittal line; two small 
very narrow sclerites on rear edge, front half of 
veniral surface with large sclerite with sagittal 
line. Mcsothorax with large rectangular dots.d 
sclente with sagittal line, with numerous strong 
setae; anterior half of metathorax with dorsal 
sclente with sagittal line, posterior half with iwo 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 



117 




c d e 

FIGURE 6. Enochrus eyrensis. a, labroclypeus; b, labium; c, maxilla; d, antenna e, mandibles. 



small foot-shaped sclerites. Legs five-segmented, 
well developed, visible in dorsal view. 

Abdomen. Segments 1 to 7 similar in shape and 
size, subdivided into three or four transverse 
folds; segment 1 with two dorsal pairs of small 
suboval sclerites, the anterior pair much the 
smaller. Segment 8 with a large, suboval, dorsal 
sclerite. Segment 9 trilobed, with a pair of short 
one-segmented urogomphi. Segments 2 to 7 with 
ventral pair of small protuberances covered with 
relatively strong, hooked setae (prolegs). 
Integument rather sparsely covered with small 
setae. 

Spiracles. Nine pairs; one mesothoracic and 
eight abdominal. Mesothoracic and first seven 
abdominal spiracles non-functional. Spiracles on 



segment 8 annular, large and functional, within 
the spiracular atrium. 

Intraspecific variation 

No significant variation within the few 
specimens seen. 

Interspecific variation 

Within the few specimens available, which 
must include at least two additional species, there 
is some variation in the distribution of setae on 
the thoracic sclerites, in the number and relative 
size of the nasale teeth, and in the length of the 
ligula. 

Specimens of E. mastersi differ more 
substantially in: pattern of setae on integument, 



'IS 



Cm WATTS 



the presence of abdominal sclerites, lack of 
prologs, the number of mandibular teeth, and Ihe 
smaller and more distal lateral protuberance on 
Ihe second antennal segment The species is 
separaiely described below. 

Identification 
By rearing larvae collected in the field. 

Remarks 

The larvae of the Australian E. (Methxdmsi 
maculictps (MacLeay) (Anderson 1976) and E. 
(M.) eyrensis differ little from those of the North 
American £. (Lumetus) ochraceus (Melsheimer) 
(Archangelsky 1997) and E (L.I fimbriatus 
(Richmond 1920, as E. perplexus tie Conte)) 
except possibly in a squatter mentum. Enochrus 
(Hydatotrephis) mastersi differs from all of the 
above by the nearly symmetrical mandibles and the 
lack of obvious sefiferous prolegs, and from £. (M.J 
eyrensis, by the more elongate and spinosc 
mentum. In these characters, other than the absence 
of prolegs, ii more closely resembles larvae of the 
subgenus Lumetus than those of Methydms. 

All known larvae of Enochrus share with most 
Helochares the angled nasale. Apart from E. 
mastersi, they are readily separated from 
tltlocharey by the asymmetrical mandibles. 
Enochrus mastersi can be separated from 
Helochares by having the Itgula shorter rather 
than longer than the basal segment of the labial 
palpus, in the lack of a basaJ patch of strong setae 
on the maxillary stipe, and in the presence of 
small dorsal sclerites on the abdominal segments. 

Specimens examined 
South Australia: 2 km S Penola. 10/99 
EihhJi'us sp Queensland. Bohle River, 10 km 

N Townsville, 23/3/96; 30 km SE Townsville 

4/5/98. South Australia: Mandina Lakes, 15/10/00: 

Tea Tree Gully, 7/5/97. 



Enochrus masters! (W MacLeay). Fig 7 

Size of third inslar. Length 12.0 mm; head 
capsule I 30 mm long, 113 mm wide. 

Head capsule, Subquadrate. Labroclypeus 
weakly asymmetrical; nasale obliquely truncate, 
left side shorter than right, with seven to eight 
short teeth, first two on right side and last one on 
left side largest; left lobe of cpislomc more 
rounded than right, shorter lhan nasale Frontal 
sulci as an inverted bell, fusing before reaching 
occipital foramen. Coronal sulcus very short. 



Gular sclerite absent. Cervical sclerites small, 
subrectangular. 

Antennae Three-segmented First segment 
wider than second segment, shorter lhan second in 
First instar larvae, about 1.5 times as long as 
second in third instar larvae, slightly bulging on 
inner apical corner; third segment about a third 
the length of the second. Sensory appendage on 
second segment much shorter than third segment. 

Mandibles. Weakly asymmetrical; Tight 
mandible with I wo strong inner teedi on basal 
half; lelt mandible with one strong inner tooth 
and one much weaker tooth. Inner margins of 
distal teeth and distal parts of mandibles serrated. 

Maxillae, Five-segmented. Stipes wide, much 
longer than remaining segments combined, with 
four strong setae on inner margin. Palpus four- 
segmented; first segment subrectangular, as wide 
as long, with an inner process a little shorter than 
second palpal segment; second segment short; 
third and four segments slightly longer, subequal 
in length 

Labium. Mentum rectangular, dorsal Surface 
with scattered culicular spines and several strong 
spines at front and side edges. Prementum 
rounded, a little wider at base, shorter than 
mentum. Palpus two-segmented, first segment 
short, second segment i.0-1 5 times longer than 
First, some small spines at the bases of both 
segments. LigttU slender, as long, as First palpal 
segment. 

Thorax and legs Prorhorax completely covered 
by a dorsal selente, with sagittal line, with 
numerous relatively long setae; ventral surface 
with two subrectangular sclerites. Mesothorax 
with a pair of large dorsal sclerites with irregular 
posterior margins, covered with relatively long 
setae. Metaihorax with narrow pair of dorsal 
sclerites each with a small foot shaped backward 
extension, covered with relatively long setae. L.ig> 
Five-segmented, well developed, visible in dorsal 
view. 

Abdomen. Segments I to 7 similar in shape and 
size, subdivided into three or four transverse 
folds; segments I (0 6 with a dorsal pair of small 
suboval sclerites. Segment 8 with a large, suboval, 
dorsal sclerite. Segment 9 irilobed, with a pair of 
short one-segmented urogomphi Integument 
covered with dense, small setae, in many pianos 
organised into tight bundles. 

Spiracles. Nine pairs; one mesothoracic and 
eight abdominal. Mesothoracic and firsl seven 
abdominal spiracles non-functional. Spiracles on 
segment 8 annular, large and functional, within 
the spiracular atrium. 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 



119 




c d e 

FIGURE 7. Enochrus masiersi. a, labroclypeus; b, labium; c, maxilla; d, antenna; e, mandibles 



fntraspecific variation 

Only three specimens are known. They vary 
slightly in size of the abdominal sclerites but 
otherwise are very similar. 

Interspecific variation 
See under E. eyrensis. 

Identification 

By association and elimination. 

Specimens examined 

New South Wales: Salisbury, 26/11/95. 
Northern Territory: Ormiston Gorge, 27/1/99. 
Victoria: King Parrot Creek, 2/12/98. 



Helochares Mulsant 

Helochares tristis W. MacLeay. Fig. 8 

Size of third instar. Length 6.8-10.0 mm; head 
capsule, 0.53-0.56 mm long, 0.66-0.69 mm wide. 

Head capsule. Subquadrate. Labroclypeus 
asymmetrical; nasale obliquely truncate, shorter 
on left side, with six small teeth; lateral lobes of 
epistome project nearly as far as nasale. Frontal 
sulci inversely bell-shaped, meeting before 
reaching occipital foramen. Coronal sulcus very 
short. Gular sclerite absent. Cervical sclerites 
naiTow, subrectangular. 

Antennae. Three-segmented. First segment as 



2fl 



a IS WATTS 



long as second dim instar larvae) or longer 
(second and third instar larvae); second segment 
with a slim apical sensory appendage on inside 
which is halt the length of third segment: third 
segment much smaller than second 

Mandibles. Slightly asymmetrical; two inner 
teeth in middle, distal one larger on left mandible, 
only slightly larger on right mandible. Distal inner 
margin and inner margins of teeth slightly 
serrated. Distal part of teeth darker. 

Miuilloe. Five-segmented. Stipes wide, longer 
than remaining segments combined, with a row of 
four stout setae on inner margin, base with a tuft 
of strong spines. Palpus four-segmented; first 
segment subrectangular, with an inner process a 
little longer than the second palpal segment; 
second segment the shortest; third segment 
longest, last segment slightly shorter than third. 

Labium. Menium large, subquadrate, with 
dorsal surface covered by strong cuticular spines. 
Premeiuuni subrectangular, wider than long. 
Palpus two-segmented, basal segment very short, 
distal segment Ihree to four limes longer than 
basal; a few small spines at base of apical 
segment. Ligula about twice as long as first palpal 
segment 

Thorax and tegs. Prothorax almost completely 
covered by a dorsal sclerite, with sagittal line; 
ventral surface with a subrectangular plate, with a 
sagittal line. Mesothorax with a pair of large 
rectangular dorsal sclerites, mctathorax with a pair 
of small dorsal sclerites, irregular in shape, 
composed of a wide and narrow basal portion and 
act L-shaped central portion arising from Ihc 
middle of the basal piece. Legs five-segmented, 
well developed, visible in dorsal view, 

Abdomen. Segment I with two dorsal pairs of 
small, narrow, irregularly shaped sclerites; anterior 
pair much smaller. Segments I to 7 similar in size 
and shape subdivided by three or four transverse 
folds; pleura of segments I to 7 with three 
limgitudinal slight bulges, the most ventral one in 
three iobes, each segment with a dorsal pair of 
setae towards the middle arising from a small 
circular sclerite or dark-pigmented area, and a 
small seta a little inwards and backwards from 
each spiracle. Segment 8 with a large, suboval, 
dorsal sclerite, serrate on posterior edge and pair 
of short apical flaps. Segment 9 trilobed; wilh a 
pair of small one -segmented urogomphi; central 
lobe largest. Integument densely covered in short 
line setae. 

SpirQCieS Nine pairs; one mesothoracic and 
eight abdominal, raised slightly above Ihe surface. 
Mesothoracic and firsi seven abdominal spiracles 



non-functional. Spiracles on segment 8 annular, 
large and functional, within the spiracular atrium 

fntraspecific variation 

There is some variation in the shape or the 
thoracic sclerites and in the shape and 
configuration of the nasalc teeth. 

Inlet specific variation 

H. luridus (W. MacLcay) and H. clypeatus 
Watts. As for H. tnstt.s. 

H. tenut.struitus Regimbait. The integument is 
predominantly covered by long thin setae rather 
than die short and very curved setae found in H. 
tristis, H. luridus and //. clyptaius. This gives a 
distinct furry look, to the larvae. 

H fdViicoffis (Montrou/.ier). Differs in lacking 
nasale, having a slight bulge on the inside nf 
antenna and a number of other characters (see 
below). These differences are sufficient to wanranl 
a separate description (sec below). 

Other Helochares species Within specimens 
unidentified to species there is variation in: form 
of ihe small setae on the integument; the shape of 
the metasternal sclerites; and the presence/ 
absence, position and shape of the abdominal 
sclerites. 

Identification 

H tristis, H luridus and H. clypeatus by rearing 
larvae collected in the field; // tenuUtriatta by 
association and elimination. 

Remarks 

Wuhin the known Australian Helochares larvae 
there arc clearly two groups: H. foveicollis: and 
H. tristis. If. luridus. H renuistriaius and H. 
clypeatus. These correspond to the subgenera 
Helochares and Hydrobaticus Blackburn, 
respectively, and strongly reinforce tire 
distinctiveness of these groups as reflected in the 
classification. Wilhtn Australian /?. 
(Hydmbaticus) there is very little difference 
between the known species or among the large 
number of unidentified larvae. In addition, there 
is minimal difference between these Australian H. 
(ifvdrobaticHS) and the North American H. 
(Hydrobaticus) maculicotlis Mulsant 
(Archangelsky l c J97: Richmond 1920) or the 
South American //. (Sindatus) talarum Fernandez) 
and H. (Helochares) pullipes (Brullc) (Feniaiidi/ 
1983) or the European H. (Helochares) obstuna 
(Mullcr) (as gtiseus Hccr) (Boving & Henrik.seii 
1938). 

The larvae of //. (Helochares) foveicollis differ 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 



121 









FIGURE 8. Helochares tristis. a. labroclypeus; b, labium; c, maxilla; d, antenna e, mandibles. 



considerably from all of these species in a number 
of characters, most noticeably the symmetrical 
labroclypeus and the more elongate antennae and 
maxillary palpi (see later). 

The larva of the related Australian genus, 
Chasmogenus, is only known from the work of 
Anderson (1976). It also has a symmetric 
labroclypeus but differs from both H. 
(Helochares) fovekollis and H. (Hydrobaticus) 
tristis in the asymmetric mandibles, one with one 
tooth and the other with two, and in the complete 
rather than partial dorsal sclerites on the 
metalhorax. 

Species of H. (Hydrabaticus) appear to breed 
more or less continuously and females are 
frequently caught with egg masses attached to 



their abdomens, although Anderson (1976) found 
that there was a distinct spring breeding season in 
H. tristis near Sydney. Larvae are among the most 
commonly collected Hydrophilid larvae, often 
found together with those of Limnoxenus in the 
south and both Limnoxenus and Sternolophus in 
the north. They can be readily separated from both 
of these by their angled nasale and mandibles with 
two rather than three teeth. 

Although adults of Enochrus are often more 
common than those of Helochares, their larvae 
are, for unknown reasons, rarely found. They 
share with H. (Hydatotrephis) the angled nasale 
but, other than E. mastersi, have strongly 
asymmetrical mandibles. Larvae of E. mastersi 
resemble those of H. (Hydatotrephis) quite closely 



122 



CHS WATTS 



but differ in lacking a cluster of strong setae at the 
base of the maxillary palpus, and in having the 
ligula shorter than the basal segment of the labial 
palpus. 

Specimens examined 

New South Wales: 20 km W Nelligan, 
3/11/97. South Australia: 10 km N Coonawarra, 
26/9/98; Cudlee Creek, 10/11/96; Kuitpo, 5/10/95; 
13 km W Meadows, 26/9/96; Mt Crawford Stale 
Forest, 10/11/96; Tea Tree Gully, 7/5/97. 
Victoria: 10 km W Cowwarr, 30/11/98; 
Healsville, 12/68. 

H. tenuistriatus. Western Australia: 30 km N 
Perth, 14/10/96 

H. clypeatus. Northern Territory: Nourlangie 
Creek, 20 km SSW Jabiru, 1 1/10/98. 

H. luridus. Queensland: Burdekin River E of 
Charters Towers, 4/5/98. 

Helocharus sp. New South Wales: 1 2 km E 
Tamworth, 26/11/95. Northern Territory: 
Nawurlandja, Kakadu National Park, 22/3/98. 
Queensland: Alligator River 20 km S Townsville, 
25/3/96; Bluewater, 22/3/96; Bowling Green Bay 
National Park, 6/11/95; 10 km N Cairns, 4/1/97; 
Eubenangee Swamp, 4/2/97; 8 km S Greenvale, 
27/3/96; Jourama Falls near Townsville, 3 1/10/95; 
1 km W Mingela, 4/5/98; Star River, 1/11/95: 
30 km SE Townsville, 4/5/98. 



H eloc hares foveicollis (Montrouzier). Fig. 9 

Size of third instar. Length 13.5 mm; head 
capsule 0.78 mm long, 0.94 mm wide. 

Head capsule. Subquadrate. Labroclypeus 
symmetrical, without nasale; with eight to nine 
small teeth along front edge in middle; lateral 
lobes of epistome slight. Frontal sulci inversely 
bell-shaped, meeting before reaching occipital 
foramen. Coronal sulcus short. Gular sclerite 
absent. Cervical sclerites small, subrectangular. 

Antennae. Three-segmented. First segment as 
long as second (first instar larvae) or longer 
(second and third instar larvae), with distinct 
protruberance on inside near apex; second 
segment with a slim apical sensory appendage on 
inside, half the length of third segment; third 
segment much thinner than second. 

Mandibles. Relatively slim, symmetrical; two 
inner teeth in middle, distal one much larger. 
Outer margin of apical tooth weakly serrated. 

Maxillae. Five-segmented. Stipes relatively 
narrow, much longer than remaining segments 
combined, with a row of five stout setae on inner 



margin, base with a few short, strong spines. 
Palpus four-segmented; first segment 
subrectangular with an inner process as long as 
the second palpal segment; second segment the 
shortest; third segment the longest; fourth segment 
shorter than third. 

Labium. Mentum large, subquadrate, with 
central portion of dorsal surface covered by weak 
cuticular spines and a semicircle of six spines, 
anterolateral angles sharp, each with a small spine. 
Prementum elongate-rectangular, longer than 
wide. Palpus two-segmented, basal very short, 
distal segment four to five times longer than basal 
one. Ligula short, thick, about twice the length of 
first palpal segment. 

Thorax and legs Prothorax almost completely 
covered by a dorsal sclerite, with sagittal line; 
ventral surface with a subrectangular sclerite, 
subdivided by a sagittal line. Mesothorax with a 
pair of large dorsal sclerites; metathorax with a 
pair of small dorsal sclerites composed of a wide 
and narrow basal portion and an L-shaped portion 
arising from the middle of the basal piece, 'heel' 
reaching posterior margin. Legs five-segmented, 
well developed, visible in dorsal view. 

Abdomen. Segments 1 to 7 similar in size and 
shape, subdivided by three or four transverse 
folds; segment 1 with two dorsal pairs of small 
narrow basal sclerites, the anterior pair smallest; 
pleura of segments 1 to 7 each with three to four 
weak longitudinal bulges, a well separated pair of 
strong dorsal setae arising from a very small 
circular darkly pigmented area, a moderately 
strong seta just inwards and behind each spiracle. 
Segment 8 with a large, suboval, dorsal sclerite 
with a ragged hind edge, and a pair of prominent 
apical flaps; segment 9 trilobed, central lobe 
largest and sclerotised, with a pair of prominent 
one-segmented urogomphi. Integument covered 
with moderately dense hair-like setae. 

Spiracles. Nine pairs; one mesothoracic and 
eight abdominal. Mesothoracic and first seven 
abdominal spiracles very small, non-functional. 
Spiracles on segment 8 annular, large and 
functional, within the spiracular atrium. 

Intraspecific variation 

Among the few specimens known there is some 
variation in the number of larger spines on the 
mentum and in wear on the nasale teeth. 

Interspecific variation 

Helochares foveicollis differs from other known 
Australian Helochares larvae by the lack of a 
nasale, presence of a prominent bulge on the 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 



123 




FIGURE 9. Helochares foveicollis. a, habitus; b, labroclypeus; c, labium; d, maxilla; e, antenna; f, mandibles. 



inside of the antennae, proportionally longer and Remarks 

narrower prementum and ligula, more elongate On adull characters H. foveicollis is presently 

maxillary palpi and the shape of the mctathoracic placed in the subgenus Helochares together with 

sclcritcs (Fig. 9). H. obscurus and H. pallipes. However larval 

characters do not support this placement, nor its 

Identification placement in either of the subgenera 

By rearing field-caught larvae and rearing from Hydatotrephis or Sindolus. 
egg mass attached to female. 



124 



CHS WATTS 



Specimens examined 

Northern Territory: Manton Dam, 2/2/99; 
2 km S Adelaide River, 1/2/99. Queensland: 
40 km S Townsville, 2/2/97. 



Hybogralius Orchymont 

Hybogralius hartmeyeri (Regimbart). Figs 10, 
17b 

Size of third instar. Length 10.0 mm; head 
capsule 1.60-1.79 mm long, 1.62-2.00 mm wide 

Head capsule. Rectangular. Labroclypeus 
asymmetrical; nasale narrow, projecting well 
forward, with five teeth, the two lateral ones more 
distant than rest; lateral lobes of epistome 
rounded, projecting farther than nasale, left lobe 
often larger, with a strong comb of stout spines on 
front edge, front edge of right lobe with a few 
scattered spines. Frontal sulci U to V-shaped, 
fusing just before reaching occipital foramen. 
Coronal sulcus short. Gular sclerite absent. 
Cervical sclerites relatively small, subrectangular. 

Antennae. Three-segmented. First segment 
longer than remaining segments combined; second 
segment about 1.5 times the length of third 
segment which is very thin. Sensory appendage 
on second segment minute. 

Mandibles. Asymmetrical; right mandible with 
three inner teeth, the anterior one largest; left 
mandible with three teeth of approximately the 
same size, central one with patch of strong setae 
at base. 

Maxillae. Five-segmented. Stipes stout, longer 
than remaining segments combined, with four 
long stout setae on inner margin. Palpus four- 
segmented; first segment widest, with a short 
inner process; second segment short; third 
segment longest; fourth segment about half length 
of third. 

Labium. Mentum small, subrectangular, sides 
slightly convex, dorsal surface with small spines, 
anterolateral angles rounded with several stout 
spines. Prementum elongate, rectangular, as long 
as mentum. Palpus two-segmented, first segment 
the shortest. Ligula present, about as long as the 
first palpal segment, tip weakly bifid. 

Thorax and legs. Prothorax with large dorsal 
sclerite, with sagittal line; ventral surface with a 
large subrectangular sclerite, with sagittal line. 
Mesothorax with a pair of large triangular dorsal 
sclerites. Metathorax with a pair of semicircular 
dorsal sclerites and a more posterior pair of much 
smaller circular ones. Sclerites, particularly the 



anterior ones, covered with very short spines, 
occasionally arranged in short rows; rest of thorax 
densely covered with short fine setae with a few 
much longer setae laterally. Legs five-segmented, 
relatively short, just visible in dorsal view. 

Abdomen. Segments 1 to 8 similar in shape, 
tapering towards the caudal end; Segments 1 to 7 
subdivided into three or four transverse folds; 
sides of each segment with some slight, fleshy 
bulges, with one to two long setae. Segment 8 
with two transverse folds, without dorsal sclerite. 
Segment 9 with hind margin weakly concave. 
Below segment 9, and projecting just beyond, are 
two prominent broad lobes. Integument densely 
covered with small fine setae. 

Spiracles. Nine pairs; one mesothoracic and 
eight abdominal. Mesothoracic and first seven 
abdominal spiracles on small papillae. Spiracles 
on segment 8 (if present at all) obsolete and 
certainly non-functional. 

Pupa (Fig. 17b). Head without styli. Pronotum 
with 22 styli, 16 around margins and 6 on disc. 
Mcsonotum and metanotum each with one pair of 
styli near the midline. Abdominal segments 1 to 7 
with a row of four styli on the terga, segments 2 
to 7 with one stylus on each pleuron; segment 8 
without styli; segment 9 with a pair of long 
urogomphi. 

fntraspecific variation 

One specimen has a slight mandibular tooth 
anterior to the normal ones (Fig. 1); otherwise, 
there is little variation other than in tooth wear 
between the few known specimens. 

Identification 
By rearing from larvae collected in the field. 

Specimens examined 

Western Australia: Gooseberry Hill, 14/9/00. 

Remarks 

Hybogralius is a monospecific Australian genus 
known only from a couple of localities in the 
escarpment region of the Darling Ranges east of 
Perth, Western Australia where it lives in small 
temporary winter/spring streams. Although placed 
with Limnoxenus in the subtribe Hydrobiina of 
the tribe Hydrophilini (Hansen 1991), the larvae 
of the two genera have little in common. Several 
features of the larvae of Hybogralius stand out: 
the lack of a spiracular atrium, the strongly 
asymmetric labroclypeus, the narrow prominent 
nasale, the strongly asymmetrical mandibles, and 
the rings of small spines at the base of the 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 



125 




a d e 

FIGURE 10. Hybogralius hanmeyeri a, habitus; b, labroclypeus; c, labium; d, maxilla; e, antenna; f, mandibles. 



segments of the labial palpi. In most of these 
characters it most closely resembles Berosus. 
Apart from the lack of a spiracular atrium and the 
presence of a coronal sulcus, it also resembles the 
larvae of Oocyclus Sharp and Larcobius in the 
tribe Laccobiini. To what degree these similarities 
reflect phylogenetic relationships remains to be 
seen. 

As well as the lack of a functional apical 
spiracle, the main tracheal trunks are narrower 
than most other genera and the spiracles are not 



much larger, if at all, than those of other genera 
where they are considered non-functional. How 
the larvae manage to breathe is a puzzle. 



Hydrobiomorpha Blackburn 

Hydrobiomorpha sp. Fig. 1 1 

Size of third instar. Length 15.0 mm; head 
capsule 1.65 mm long, 1.70 mm wide. 



126 



CHS WATTS 








c d e 

FIGURE 1 1. Hydrobiomorpha sp. a, labroclypeus; b, labium; c, maxilla; d, antenna; c, mandibles 




Head capsule. Subquadrate. Labroclypeus 
slightly asymmetrical; nasale truncate, edge rough 
rather than toothed; lateral lobes of epistome 
rounded, projecting further than nasale. Frontal 
sulci V-shaped, meeting before reaching occipital 
foramen. Coronal sulcus short. Gular sclerite 
absent. Cervical sclerites small, subquadrate. 
Numerous small darker patches on posterolateral 
angles and within frontal sulci. 

Antennae. Three-segmented. First segment 
slender, three times longer than remaining 
segments combined (third instar larvae), inner 
margins with numerous short spines; second 
segment as long as third. Sensory appendage of 
second segment very small. 



Mandibles. Symmetrical, sharply pointed; right 
mandible with three teeth on basal half, distal one 
large, bifid apically, central tooth smaller, basal 
one much smaller; left mandible similar to right 
except for basal tooth placed slightly more 
ventrally. 

Maxillae. Five-segmented. Stipes slender, 
longer than remaining segments combined, with a 
group of short spines on inner margin at base, and 
several setae along inner and outer margins. 
Palpus four-segmented; first segment longest, with 
a short inner process; second segment the shortest; 
third and fourth segments subequal in length, each 
twice as long as second segment. 

Labium. Mentum approximately square, 



LARVAE OF SOME AUSTRALIAN AQUATIC UYDROPHILIDAE 



127 



anterolateral angles projected, each with a pair of 
very small spines, dorsal surface with eight short, 
stout setae on distal two-thirds, and short cuticular 
spines on hasal third. Prementurn longer than 
wide, half the width of mentum. Palpus Iwo- 
segmented, distal segment three to four times 
longer than basal segment. Ligula a little longer 
than first palpal segment, weakly bifid. 

Thorax and legs- Prothorax covered hy a large 
dorsal selcrile, posterior end rounded, sagittai line 
present; ventral sclente large, subrectangular, with 
sagittal line. Mesothorax with pair of irregularly 
shaped sclentes, wider anteriorly, with sagittal 
line. Metathorax with pair of small irregular 
shaped seleriies. Legs five-segmented, long, 
visible in dorsal view. Prothorax and mesothorax 
covered with small, scattered, dark patches. 

Abdomen . First segment subdivided into two 
IranSverse folds, with a dorsal pair of narrow 
sclerites near anterior edge; segments 2 to 7 
subdivided into four or five transverse folds, 
without sclerites. Segments 1 to 7 each with 
eight setose tubercles, four dorsal and two on 
each lateral margin; pleura weakly lobed. 
Segment 8 with a small, suboval, dorsal sclcrite, 
posterior apex subdivided into four Johcs and a 
pair of small procerei. Segment 9 round, with 
three small dorsal sclerites, bearing a pair of 
short, one-segmented urogomphi, and a pair of 
long paracerci, a pair of gill-like appendages 
(prostyli) originate on ventral side Integument 
sparsely covered with relatively long setae. 
Single, narrow, darker line along midline oi 
dorsal surface. 

Spiracles Nine pairs; one mesothoiaeic and 
eight abdominal. Mesothoracic and first seven 
abdominal spiracles non- functional Spiracles on 
segment 8 annular, large and functional, within 
the spiracular atrium 

Identification 

Agreement with generic description by 
Archangel sky (1997). Specimens were tuken in 
association with H bovilli Blackburn and arc 
almost certainly that species. 

hitrrspecific variation 

There is little variatiun between liic two known 
specimens. 

Hemarks 

The larvae described above differ hide from 
those of the New World H. casta (Arehangelsky 
1997; Spangler 1973) or die unidentified African 
species described by Berge Henegouwcn (1982). 



The peculiar shape of ihe anterior mandibular 
tooth is diagnostic within Ihc Australian fauna. 

Spi i imens exam itnJ 
Northrrn Territory: Holmes Jungle. 28/1 1/99. 



Hydrophilus Geuftroy 

Hydrophtlus billneatm (MacLeay). Fig. 12 

Size of third instar Length 35 mm; head 
capsule 3.25 mm long. 4.25 mm wide. 

Head capsule. Suboval. Labroclypeus virtually 
symmetrical; nasalc undeveloped: lateral lobes of 
epistomc rounded, projecting farther than nasale. 
Frontal sulci broadly U-shaped, fusing just before 
reaching occipital foramen. Coronal sulcus 
present, very short. Gular sclerite absent. Cervical 
sclerites small, suboval. 

Antennae. Four-segmented. First segment 
slender, slightly constricted near base, longer than 
remaining segments combined, with some slender 
selae on distal two-thirds of inner margin, second 
segment short, constricted near base in first instar 
larvae, third and fourth segments subequal in 
length. Sensory appendage on third segment 
reduced to a small papilla. 

Mandibles. Asymmetrical; right mandible 
longer, more slender, with a large tooth on basal 
half which is divided by a longitudinal groove 
into ventral and dorsal sections; left mandible 
shorter, with one small inner tooth on basal hall 

Maxillae. Five-segmented. Stipes narrow, 
elongate, longer than remaining segments 
combined, with three stout setae on inner margin. 
Palpus four-segmented: first segment the longest, 
with a short inner process; second and third 
segments subequal in length; fourth segment 
slightly shorter. 

Labium. Menium subrectangular, sides convex, 
anterolateral angles lobed. central third with sonic 
small spines (without spines in first and second 
instars). Prementurn subtrapezoidal, anterior end 
wider with a pair of small protuberances towards 
front. Palpus two-segmented, first segment 
shortest. I.igul3 present, much Shorter lhan first 
palpal segment. 

Thorax and legs. Prothorax with two large 
sclerites together covering most of dorsal surface, 
the triangular area anterior to these scleriies 
usually sclerotised; ventral surface with a large 
subrectangular sclente, with sagittal line. 
Mesothorax with a pair of subtriangular dorsal 
sclerites. Metathorax with a pair of narrow, basal 



128 



CHS WATTS 




o o o 



a 






c d e 

FIGURE 12. Hydrophilus bilineatus. a, labroclypeus; b, labium; c, maxilla; d, antenna; e, mandibles. 



sclerites and a more central pair of small 
irregularly-shaped ones. Legs five-segmented, 
relatively short, visible in dorsal view, with rows 
of long swimming setae on both dorsal and ventral 
edges of femur and tibiotarsus. 

Abdomen. Segments 1 to 6 similar in shape, 
tapering towards the caudal end; segments 7 to 8 
more elongate and slender. Segments 1 to 7 
subdivided into three transverse folds; third fold 
with two lateral and two dorsal small tubercles, 
more obvious on posterior segments. Pleura 1 to 8 
each with a small lateral tubercle. Segment 8 with 
two small subtriangular sclerites at posterior end; 
segment 9 slightly trilobed, with a pair of short 
one-segmented urogomphi, ventromedially with a 
pair of long gill-like appendages. Integument in 
first instar very densely covered with small thin 
setae; in second and third instars moderately 



covered with thin, relatively long setae and 
numerous small 'bumps'. 

Spiracles. Nine pairs; one mesothoracic and 
eight abdominal. Mesothoracic and first seven 
abdominal spiracles very small, non-functional. 
Spiracles on segment 8 annular, large and 
functional, within the spiracular atrium. 

Identification 
By association and elimination. 

Intraspecific variation 

I have not noticed any significant variation 
within the few available specimens I have 
identified as H. bilineatus. 

Interspecific variation 
Within the larvae available there is considerable 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 



i y. 



variation in a number of characters, most notably: 
number of antennal segments vary from three lo 
four, the prescncc/abscnce/form of the lateral 
piojcctitms on '.he abdominal segments; form uf 
covering of the integument; dorsal colour pattern. 

Remarks 

The larvae of two European species, H piceus 
Linnaeus and //. aterrimus Eschscholtz. and the 
North American //. triangularis (Say) arc well 
known (Archangelsky 1907: Boving & 
llennksen 193K: Richmond 1920). In addition, 
the larvae of // senegalensis (Pcrcheron) (flerge 
Henegouwcn 1982; Africa) and H. acuminatus 
Motschulsky (Morioka 1955; Japan) have heen 
described. Within the known larvae there is 
considerable variation in the number of antennal 
segments in different instars. in the development 
of lateral abdominal flaps and, at least within the 
Australian species, in the form of the integument 
coverage. Somewhat unusually for Hydrophilidae 
latvae, it appears that specific differences are 
sufficient to enable the larvae of many species to 
be identified. 

In the lack of lateral abdominal flaps and its 
Unit segmented antennae, H. bilineaius most 
closely resembles the North American H. 
triangularis but differs in a number of details, 
most obviously in the more rohust mandibles with 
a well-developed molar region. The other 
Australian species appear more distant, 
particularly in the well-developed abdominal Haps 
and number of antennal segments. With the more 
robust mandibles and abdominal flaps these 
Australian species appear to be closer to Ihe 
subgenus Dihhcelux (sensu Hansen 1991) than 
subgenus Hydmphilus i Archangelsky 1997). On 
adult characters they fit clearly into the subgenus 
Hxdrophilus. 

Egg cases of several species were collected in 
the Held and each produced between 15 and 30 
larvae. Those thought to be of H bilineatu.s 
refused all food offered but the other species 
accepted, often with some excitement, small snails 
and some accepted freshly killed mosquito larvae. 
None accepted chironomid larvae. Unfortunately, 
despite seemingly accepting snails as food, none 
was reared to the second instar. Species varied in 
the degree of cannibalism. Some broods were 
quite strongly cannibalistic buc in others, apart 
from the occasional unfortunate individual, 
siblings lived well together. 

Specimens of Hydmphilus bilimatus 
iMacLeay) have previously been identified as II. 
piiicarius Chevrolat t Hansen 1999i. 



Specimens examined 

Northern Territory: Manlon Dam, 23/3AJ7; 
2 km S Adelaide River 1/2/99 

Hydrophilus sp. Northern Territory: Newry 
Station, 2/S6 col. M. J Tvler Queensland: 25 km 
S Townsville, 3/5/98, 2/2/97; 8 km S Geenvale 
27/3/96; 5 km NE Mt Molloy. 30/3/96. South 
Australia 10 km N Coonawarra, 10/11/97, 
16/10/97; 1 km S Nangwarry, 9/10/97. Tasmania: 
12 km N Hobarl, 2/12/00; 2 km W Finglc, 
23/1/00; 3 km SW Clifton Beach, 4/12/00. 
Victoria: 5 km NE Dartmoor. 1 1/10/97 Western 
Australia: Murchison River, 13/3/95, col. S. 
Hal.se: 6 km S Pinjarrah, 23/10/96. 

Laccobius Erichson 

Laccobim decipiens Gentili Fig. 13 

Size of third instar. Length 5.0-6.5 mm; head 
capsule 0.43-0.50 mm long. 0.45-0.46 mm wide. 

Head capsule. Subrectangular. Labroclypeus 
strongly asymmetrical; nasale prominent, with 
three teeth, lateral lobes of epistome 
asymmetrical, both projecting further than nasale; 
left lobe larger, covering basal third of mandible, 
with a row of about 12 strong curved setae un 
inner margin, larger ones dentate; right lobe 
smaller, covering a quarter of mandible, withouc 
setae. Frontal sulci parallel, reaching occipital 
foramen without coming together, difficult to see 
in thud instar larvae. Coronal sulcus absent. Gular 
scleritc absent; cervical sclentes small and oval, 
difficult to .sec in first instar larva. 

Antenna. Thrcc-scgmcntcd. second segment the 
longest, with a small outer sensory appendage 
about a quarter the length of third segment; first 
segment half the length (first instar larvae) to 
slightly shorter (third instar latvae) than second 
segment. 

Mandibles. Strongly asymmetrical. Left 
mandible with three inner teeth, central one 
largest, with five strong spines, region behind 
third tooth with a number of very short spines: 
right mandible with two inner teeth, larger than 
those of left mandible, front one much larger, 
occasionally also with a very small third tooth, or 
spine, behind the second. 

Maxillae. Five-segmented. Stipes wide, longer 
than remaining segments combined, inner margin 
with four setae. Palpus four-segmented: first segment 
subquadrate, incompletely sclerotiscd, with a small 
inner process; second segment shortest, third and 
fourth segments subequal in length. 



130 



CHS WATTS 




a d e f 

FIGURE 13. Laccobius decipiens. a, habitus; b, labroclypeus; c, labium; d, maxilla; e, antenna; f, mandibles. 



Labium. Mentum small and narrow, 
subrectangular. Prementum slightly larger, 
trapezoidal. Palpus two-segmented, first segment 
shorter, second segment two to three times longer 
than first. Ligula obsolete, at most reduced to a 
slight bulge. 

Thorax and legs. Prothorax completely covered 
by a large dorsal sclerite, with sagittal line; 
ventrally with a pair of large subrectangular 
sclerites, fused in anterior half. Mesothorax with 
two pairs of dorsal sclerites, anterior one small, 
subrectangular; posterior pair larger, 
subtriangular. Metathorax with one pair of small, 
irregularly shaped sclerites intermediate in size to 
the two mesothoracic ones. Legs five-segmented, 
visible in dorsal view. 



Abdomen. Segments 1 to 7 similar in shape, 
tapering towards posterior end, "segments 
subdivided by transverse folds; sides with several 
slight bulges, each segment with a pair of long 
dorsal setae arising from small, darkly pigmented 
areas, one just behind spiracle and the other near 
midline. Segment 8 with a subtriangular dorsal 
sclerite. Segment 9 small, trilobed, bearing a pair 
of small one-segmented urogomphi. Integument 
with quite dense covering of short fine setae. 

Spiracles. Nine pairs; one mesothoracic and 
eight abdominal. Mesothoracic and first seven 
abdominal spiracles non-functional, on short dark- 
pigmented papillae; spiracles on segment 8 
annular, large and functional, within the spiracular 
atrium. 



LARVAE. OF SOME AUSTRALIAN AQUATIC HYDROPHIUDAE 



131 



Intraspecific variation 

Wiihin specimens that I have identified as this 
species theie is some variation as foiiows: some 
specimens have an additional small third tooth on 
Ihe right mandible, in newly moulted specimens 
there are slight serrations on the upper mandibular 
leeth which appear to wear away quickly; (he first 
abdominal segment occasionally has a pair of very 
small dorsal sclcrites. 

Interspecific variation 

Within specimens unidentified to species there 
is Variation in the Shape of the dorsal sclerites on 
abdominal segment 8; the width of the nasale and 
the shape of the nasale teeth; the presence, size 
and position of abdominal sclerites. Overall these 
differences are slight. 

Identification 
By rearing larvae collected in the field. 

Specimens examined 

New South Wales. Bomhala, 28/11/98; ditto, 
4/11/97; ditto. 18/1/97. Victoria; Stratford, 
7/1 1/97. 

Liccobius spp Queensland: Kauri Creek near 
Tinarro Dam, 24/10/9?, col D Larson. 

Remarks 

There is little difference between the larvqe of 
the Australian species and those of the North 
American L minuwides Orchymont and L. agilis 
(Randall) described and illustrated by Richmond 
( 1 920) and Archangelsky ( 1 997 ). 



Limnoxenus Motschulsky 
Limnoxenus zeolandicus (Broun). Fig 14 

Size of third instar. Length 1 1 5-15,0 mm; head 
capsule 1.60-1.79 mm long. 1.62-2.00 mm wide. 

Head capsule Square. Lahroclypcus slightly 
asymmetrical; nasale moderately developed, 
angled slightly with right side more foiward, with 
live teeth, left tooth a little distant from others; 
lateral lobes of epistorrte rounded, projecting not 
quite as far as nasale, right lobe often a litile more 
rounded and a little inure projected than left lobe, 
Frontal sulci U to V-shaped, fusing just before 
reaching occipital foramen. Coronal sulcus very 
short Gular sclerite absent. Cervical sclerites 
relatively small, subrectangular. 

Antennae, Three-segmented First segment 
longer than remaining segments combined, with 



some slender setae on distal two-thirds of inner 
margin; second segment about twice the length of 
third segment which is very Ihin Sensory 
appendage on second segment minute, 

Mandibles. Symmetrical with three inner teeth. 
progressively smaller towards base, 

Maxillae. Five-segmented. Stipes stout, longer 
than remaining segments combined, with four 
long stout setae on inner margin. Palpus four- 
segmented; first segment the widest, with a short 
inner process: second and fourth segments short, 
subequal in length: third segment longest. 

Labium. Mentum subteetangular, sides slightly 
concave, dorsal surface with small spines, 
anterolateral angles rounded, with several stout 
spines. Prementum square, not much smaller than 
mentum. Palpus two- segmented, first segment the 
shortest. Ligula present, slightly longer than first 
palpal segment, tip bifid 

Thorax and legs. Prnthorax with large dorsal 
sclerite, with sagittal line; ventral surface with a 
large subrcctangular sclerite, with sagittal line. 
Mesothorax with a pair of large triangular dorsal 
sclerites. Metathorax with a pair of wineglass- 
shaped dorsal sclerites sometimes with stem of 
wineglass absent. Rest of surface of thorax 
covered with very short spines, occasionally 
arranged in short rows. Legs five-segmented, 
relatively short, barely visible in dorsal view. 

Abdomen. First segment with a dorsal pair ol 
small subovoid sclerites close to anterior margin. 
Segments I to 6 similar in shape, tapering towards 
the caudal end; segments 7 and 8 more elongate 
and slender Segments 1 to 7 subdivided into three 
transverse folds; dorsal surface of third fold with 
four small, dark, narrowly cylindrical tubercles: 
sides of each segment with three slight, fleshy 
bulges. Segment 8 with a large subnval dorsal 
sclerite, with three to four quite deep indentations 
on hind edge, lighter coloured towards middle 
Segment 9 trilobed, with a pair of short one- 
segmented urogomphi. Integument densely 
covered with small setae and a sparse covering of 
longer setae which get progressively smaller 
posteriorly. Dorsal surface often with two darker 
stripes through sclerites and inner pair of 
tubercles. 

Spiracles. Nine pairs: one mesothoracic aruJ 
eight abdominal Mesothoracic and first seven 
abdominal spiracles non-functional, on small 
papillae. Spiracles on segment 8 annular, large 
and functional, withiu the spiracular auium. 

Intraspecific variation 
The number of nasale teeth is variable with up 



132 



CHS WATTS 








a d e f 

FIGURE 14. Litnno.xenus zealandicus. a, habitus; b, labroclypeus; c. labium; d. maxilla; e, antenna; f, mandible. 



to ten smaller teeth in some specimens; in some Identification 

specimens these are worn down to the stage where By rearing from larvae collected in the field. 

the front of the nasale is almost smooth. The 

central nasale teeth are not well developed in first Remarks 

instar larvae. In one specimen the antennae are The larva of the European L. niger (Gimelin) 

four segmented with the normal third segments has already been described by Berge Henegouwen 

clearly divided into two. (1975). The larva of the sole Australian and New 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHILIDAE 



133 



Zealand species differs Jinle I'rom those of this 
species. 

One of the most commonly collected 
hydrophilid larvae in Australia, it is quickly 
recognised by the parallel darker stripes on the 
dorsal surface and combination of relatively short 
basal segment to the antenna and symmetrical 
three-toothed mandibles. The dense covering of 
small setae gives the abdomen a fur-like look. 

Specimens examined 

New South Wales: 1(1 km E Braidwood. 30/1 1/95; 
Bra id wood, 19/1/97 

South Australia. I Okm N Coonawarra, 10/11/97; 
10 km E Mt Compass, 13/9/97; Ml Crawford 
Forest, 10/11/96; 6 km N Forreslun. 3/10/95; 
19 km N Forreston, 3/9/99; 1 km S Nangwarry, 
9/10/97; ditto. 29/8/99; 2 km S Penola, 20/i 1/99; 
Warburtott River, 2-8/10/99, 27°52'23 "S 
137°54-40'E; Watervalley, 7.2 km NNE Mt 
Rough, 15/10/00. Tasmania; Lake SI Clair, 4 km 
N Dcrwent Bridge, 25/1/00 Victoria 5 km NE 
Dartmoor, 11/10/97; 2 km W Brimpaen, 23/9/98; 
Hcalsville. 12/68; 6 km N Noojee, 16/1/97 
Western Australia: Gin Gin, 15/10/96; 10 km E 
Kalamunda, 16/10/96, 2 km W Nannup, 20/10/96; 
6 km S Pinjarrah, 23/10/96; 1 km S Serpentine, 
24/10/96. 10 km S Yallingup, 22/10/96. 



Regimbartia Zaitzev 

Rtgimbariia attenuata (Fabncius), Fig, 15 

Size of third instcir. Length 5.0-7.5 mm; head 
capsule 1.00-1.0] mm wide, 0.75-0 80 mm long. 

Head capsule. Subuuadrangular Labroclypeus 
symmetrical; nasale very short, with numerous 
short teeth on anterior border of epistome; lateral 
lobes of epistome rounded, very short, not 
projecting as far as nasale. each with two long 
setae. Frontal sulci straight, short, meeting at 
about the level of the antcnnal bases. Coronal 
sulcus long, about half the length of the head 
capsule. Gular sclvnte apserlt. Cervical sclcrites 
small, subrectangular 

Antennae. Three-segmented. Hirst segment 
slender, longer than other two combined, with a 
prominent subapical inner process some distance 
from apex. Sensory appendage on second segment 
slim, slightly shorter than third antennal segment. 

Mandibles. Virtually symmetrical, long and 
slender, with two inner teeth on basal half; distal 
tooth large, basal one small, 

Maxillae. Five-scgmenled. Stipes slender, much 



longer than remaining segments combined, wilh 
four or five setae on inner margin. Palpus four- 
segmented; third segment longest, second segment 
shortest, first and fourth segments subequal in 
length, first segment with a short inner process. 

iMbium. Menlum large, wider than long, sides 
convex, dorsal surface with cuticular spines 
Pretnentum elongate, much narrower than 
mentiim. Palpus two-segmented, basal segment 
short. Ligula annular, two to three limes as long 
as basal segment of palpus. 

Thorax and le>>s. Prothorax with a large dorsal 
sclcrilc, with sagittal line; surface covered by 
minute spines; fringe ol long, slender setae around 
the margins, four small spines, one on each 
anterolateral angle and two towards the middle; 
ventral surface with large, subrectangular sclcritc. 
without sagittal line. Mesothorax with two dorsal 
pairs of subtrtangular sclerites; metathorax with 
one dorsal pair of irregular shaped sclerites. Both 
mesothorax and metathorax with five pairs of 
seliferous projections, about half of (hem wilh a 
narrow cylindrical projection with a long seta 
attached at the end. Legs five-segmented, long., 
visible from above. 

Abdomen. Segments 1 to 7 with five pairs of 
setifcrous projections similar to those on thorax, 
four lateral ones with one more posterior than 
others and one closer to midline. In addition, Hie 
abdomen is covered with small stellate setae and 
the dorsal surface wilh small projections each 
with six to eight long, sharply pointed setae (in 
most specimens these structures are obscured by 
an accumulation of sand grains and detritus) 
Segment 8 without setiferous projections, with 9 
pair of small linger-like 'proceref, with dotsal 
semispherical sclente which covers the spiraeulai 
atrium. Segment 9 trilobed, with a pair of small, 
unsegmented urogomphi. 

Spiracles. Nine pairs; one mesothoracic arid 
eight abdominal Mesothoracic and first seven 
abdominal spiracles non-functional. Spiracles on 
segment 8 annular, large and functional, within 
the spiracular atrium. 

Interspecific variation 

There is little variation between the few 
specimens seen. 

Identification 

By association of adults and larvae by 
isoenzyme electrophoresis. 

Remarks 
The larvae closely resemble both AUocotocerus 



134 



CHS WATTS 




FIGURE 15. Regimbartia attenuata. a, habitus; b, labroclypeus; c, labium; d, detail of abdominal Stella; e, tip of 
antenna; f, maxilla; g, antenna; h, mandibles. 



and the New World Derallus (Archangelsky 1997; 
Spangler 1966), all three genera showing a suite 
of unusual characters that clearly separate them 
from other Hydrophilid larvae. The differences 
between the genera are relatively slight (see key 
and under Allocotocerus). 

Specimens examined 

Queensland: 6 km N Bluewater, 3/2/97; 15 km 
W Mareeba, 6/12/90, col. D. Larson; 5 km NW 
Mareeba, 22/9/90, col. D. Larson; 2 km N Mt 
Molloy, 1/4/96. 



Sternolophus Solier 

Sternolophus marginicollis (Hope). Fig. 16 

Size of third instar. Length 11.5-15.0 mm; head 
capsule 1.80-1.90 mm long, 1.55-1.70 mm wide. 

Head capsule. Subrectangular. Labroclypeus 
weakly asymmetrical; nasale short, weakly 
projecting with five short teeth, left one a little 
distant from rest; lateral lobes of epistome 
symmetrical not projecting beyond nasale, 
undulating. Frontal sulci V-shaped, fusing before 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPHIL1DAE 



135 





a d e f 

FIGURE 16. Sternolophus sp. a, habiius; b. labroclypeus; c, labium, d. maxilla; c, antenna; f. mandible. 



reaching occipital foramen. Coronal sulcus short. 
Gular sclerile absent. Cervical sclerites small, 
suboval, longitudinally oriented. Frontal sulcus 
outlined in a darker colour. 

Antennae. Three-segmented. First segment 
much longer than other two combined, with an 
annular ring of long setae near apex in second and 
third instars; second segment and third segment 
subequal. External apical appendage on second 
segment lacking; a small flat button- like structure 
on outside towards apex. 

Mandibles. Virtually symmetrical, with three 
inner teeth, distal two large, basal one much 
smaller. 

Maxillae. Five-segmented. Stipes as long (first 
instars) or longer (second and third instars) than 



remaining segments combined, with four or five 
setae on inner margin and patch of small spines 
on inside near base. Palpus four-segmented; first 
segment with an inner apical process; second 
segment short; third and fourth segments a little 
shorter than first. 

Ixtbium. Mentum with sides convex, slightly 
wider than long, anterolateral corners pointed, 
dorsal surface with numerous small cuticular 
spines and row of longer spines towards apex; 
small spines mostly absent in first tnstar. 
Premcntum long and thin. Palpus two-segmented, 
basal segment much shorter than distal. Ligula a 
little shorter than distal segment of palpus, slightly 
bifid at tip. 

Thorax and legs. Prothorax with large dorsal 



136 



CHS WATTS 



sclerite, with sagittal line, with pattern of darker of prothorax. Legs five-segmented, long, visible 

lines and dots; ventral surface with a large in dorsal view, with rows of long swimming-setae 

subrcctangular sclerite, with sagittal line, on femur and tibiotarsus. 

Mesothorax and metathorax each with a dorsal Abdomen. Segments I to 6 similar in shape, 

pair of subtriangular sclerites, smaller than those tapering towards the caudal end: segments 7 and 8 




FIGURE 17. a. Amphiaps queensltmdicus: ventral, lateral and dorsal views of pupa, b, Hybograliw; hartmeyeri: 
ventral and dorsal views of pupa. 



LARVAE OF SOME AUSTRALIAN AQUATIC HYDROPH1LIDAE 



137 



moie elongate and slender. Segments 1 to 7 
subdivided into three transverse folds, each 
segment with uansverse row of six pairs of long 
setae arising from shon dark-pigmenled cylindrical 
bases on dorsal surface of posterior fold. Pleura I 
to 8 each with a small, lateral, setifcrous tubercle. 
Segment 8 with two small, dorsal, subtriangular 
sclerites at posterior end. Segment 9 slightly 
inlobed, with a pair of short one-segmented 
urogomphi. Integument densely covered with both 
very' short and moderately long, setae. 

Spiracles. Nine pairs; one mesothoracic and 
eight abdominal. Mesothoracic and first seven 
abdominal spiracles non-functional. Spiracles on 
segment 8 annular, large and functional, within 
the spiracular atrium. 

Intraspecijic variation 

There is some suggestion of two longitudinal 
dark lines on the dorsal surface in some 
specimens, and in an occasional specimen the 
small third mandibular tooth is virtually absent. 

Interspecific variation 

In some specimens of Sternolophus there is a 
greater development of the lateral abdominal 
tubercles into quite prominent finger-like 
projections, especially towards the rear, a row of 
setifcrous papillae across the abdominal segments 
instead of small cylinder-like structures; a more 
symmetrical nasale; a stronger third mandibular 
tooth, and some difference in the shape of the 
meso and metathoracic sclerites (the habitus 
drawing (Fig. 16) is of this form). These larvae 
are S. centralis Watts and/or S. immarginatus 
Orchymonl. The distribution of specimens would 
suggest die lattei but not certainly so. 

Identification 
By rearing from field-collected larvae. 

He marks 
The larva of 5 marginicollis is one Of the 



few Australian larvae already known (Bertram! 
1935) as S. tenehricosus Blackburn, a junior 
synonym (Walls 1989), although the 
identification was by association, which is 
unsafe in this genus because it often has two 
or more species common ai the same locality. 
Other species have been described from Africa 
and Japan (see Berirand 1972). The present 
larvae, certainly of S-. marginicoltis and 
probably also including S. immarginatus 
Orchymonl, do nol differ significantly from 
the published descriptions and illustrations. 

Larvae of Sternolophus are commonly 
encountered in Northern Australia and can be 
readily separated from other llydrophilid larvae 
by the combination of Jong thin prementum, long 
ligula and mandibles with three teelh. 

Specimens examined 

Northern Territory: Jabiru. 19/3/98; Manton 
Dam, 2/2/99; Ormiston Gorge, 27/1/99 
Queensland: Killymoon Creek, 25 km S 
Townsville, 2/2/97. 

Sternolophus sp. Northern Territury: Holmes 
Jungle near Darwin, 28/1/99. Queensland: 8 km 
N Bluewater, 31/10/95; Bohlc River, 10 km N 
Townsville, 23/3/96; Eubcnangee Swamp, 4/2/97; 
2 km S Mt Molloy, 30/3/96. 



ACKNOWl.filXjMFNTS 

1 wish to thank Mr Rod Gultendge, Ms Elocsc 
O'Grady and Mr Harold Hamci for so ably preparing 
the illustrations; Dr David Larson for donating larvae he 
collected whilsl working in Nonh Queensland: the 
South Australian Library staff. Ms Marianne Anthony 
and Ms Jill Evans, for procuring references lhal we did 
not already have in the libniry: and Ms Debbie Churclics 
for preparing the final manuscript copy 1 would 
particularly like to thank Dr Steve Donne Man of die 
Evolutionary Biology Unit ot the Smith Australian 
Museum tor undertaking the biochemical matching of 
adult and larval Allocotocerus and Regimbartia. 



RwEBENCBS: 



Anderson, JME. 1976. Aquatic Hydrophilidac 
(Colcoptera). The Biology of some Australian 
species with descriptions of immature stages in the 
laboratory. Journal of the Australian Entomological 
Society 15: 219-228- 

Arr.hsntjelsky. M 1997 Studies on the biology, ecology 
and systematic* of die immature stages of new world 
llydrophiloidea (Colcoptera: Staphylimformia) Ohio 
Biological Survey Bulletin New Series 12: 1—207. 



Berge Henegouwen, AL van. 1975. Description of the 
egg-case and larva of Limnoxenus niger (Zschach) 
(Colcoptera Hydrophilidac) Entomologische 
Berkiucn Amsterdam 35. 27-30. 

Bergc Henegouwen. AL van. 1982. Notes on the larval 
stages of sume East African Hydiopiiiliniic 
(Colcoptera: Hydlophilidae) Entnmnlngisihe 
Hcrkhlen Amsterdam 42: 1 1-16. 



138 



CHS WATTS 



Bertrand, H. 1935. Larves de Coleopteres aquatic de 
l'Expedition Limnologique Allemande en Insulinde. 
Archiv fur Hydrobiologie 6: 193-215. 

Bertrand, H. 1972. Larves et nymphes des Coleopteres 
aquatic du globe. F. Paillart: Abbeville, France. 

Boving, AC & Henriksen, KL. 1938. The developmental 
stages of the Danish Hydrophilidae (Ins. Coleoptera). 
Videnskabelige Meddelelser fra Dansk Naturhistorik 
Forening i Khobenhaven 102: 27-162. 

Fernandez, LA. 1983. Helochares (Sindolus) talarium 
sp. no v., redescripcion de Helochares (Helochares) 
pallipes (Brulle) y descripcion de sus estados 
preimaginales. Coleoptera, Hydrophilidae. Limnobios 
2: 439^149. 

Gentili, E. 1980. The genera Laccobius and Notohydrus 
(Coleoptera, Hydrophilidae) in Australia and New 
Zealand. Records of the South Australian Museum 
18: 143-154. 

Gentili, E. 1992. The Notohydrus of Australia 
(Coleoptera, Hydrophilidae). Bolettino Society 
Entomologia Italiano, Genova 124: 21-26. 

Gentili, E. 1993. Paranacaena Blackburn, 1889: a valid 
genus (Coleoptera, Hydrophilidae). Gioruale italiano 
di entomologia 6: 285-296. 

Gentili, E. 2000. The Paracymus of Australia 
(Coleoptera: Hydrophilidae). Records of the South 
Australian Museum 33: 101-122. 

Hansen, M. 1991. The Hydrophiloid beetles. Phytogeny, 
classification and a revision of the genera 
(Coleoptera, Hydrophiloidea). Biologiske Skrifter 40: 
1-367. 

Hansen, M. 1999. Hydrophiloidea (s. str.) (Coleoptera). 
In 'World Catalogue of Insects', vol. 2: 1-416. 
Apollo Books: Stenstrup, Denmark. 

Morioka, A. 1955. Description of four Hydrophilid 
larvae from Japan (in Japanese). Shin-Konchu 8: 15- 
18. 

Oliva, A. 1992. Cuticular microstructure in some genera 
of Hydrophilidae (Coleoptera) and their phylogenetic 



significance. Bulletin de I'Institut Royal des Sciences 
Naturelles de Belgique, Entomologie 62: 33-56. 

Richmond, EA. 1920. Studies on the biology of aquatic 
Hydrophilidae. Bulletin of the American Museum of 
Natural History 42: 1-94. 

Spangler, PJ. 1966. A description of the larva of 
Derallus rudis Sharp, (Coleoptera, Hydrophilidae). 
The Coleopterists Bulletin 20: 97-100. 

Spangler, PJ. 1973. A description of the larva of 
Hydrobiomorpha casta (Coleoptera: Hydrophilidae). 
Journal of the Washington Academy of Science 63: 
160-164. 

Watts, CHS. 1963. The larvae of Australian Dytiscidae 
(Coleoptera). Transactions of the Royal Society of 
South Australia 87: 23-40. 

Watts, CHS. 1987. Revision of Australian Berosus 
Leach (Coleoptera: Hydrophylidae). Records of the 
South Australian Museum 21: 1-28. 

Watts, CHS. 1988. Revision of Australian Hydrophilus 
Muller, 1764 (Coleoptera: Hydrophilidae). Records 
of the South Australian Museum 22: 1 17-130. 

Watts, CHS. 1989. Revision of Australian Sternolophus 
Solier (Coleoptera: Hydrophilidae). Records of the 
South Australian Museum 23: 89-95. 

Watts, CHS. 1990. Revision of the Australian 
Hydrobiomorpha Blackburn (Coleoptera: 
Hydrophilidae). Records of the South Australian 
Museum 24: 35-42. 

Watts, CHS. 1995. Revision of the Australasian genera 
Agraphydrus Regimbart, Chasmogenus Sharp and 
Helochares Mulsant (Coleoptera: Hydrophilidae). 
Records of the South Australian Museum 28: 1 13— 
130. 

Watts, CHS. 1998a. Revision the Australian Amphiops 
Erichson, Allocotocerus Kraatz and Regimbartia 
Zaitzev (Coleoptera: Hydrophilidae). Records of the 
South Australian Museum 30: 93-106. 

Watts, CHS. 1998b. Revision of Australian Enochrus 
(Coleoptera: Hydrophilidae). Records of the South 
Australian Museum 30: 137-156. 



EMBOLOCEPHALUS YAMAGUCHI (BRINKHURST, 1971) (CLITELLATA: 
TUBIFICIDAE) FROM SOUTH AUSTRALIAN STREAMS 

A. M. PlNDER & P. K. MCEVOY 



Summary 

Papillate tubificids from streams in the Mount Lofty Ranges are identified as Embolocephalus 
yamaguchii (Brinkhurst, 1971) and represent the first records of this largely Holarctic genus from 
Australia. Collection details are provided for records of a second papillate species, identified as 
belonging to the genus Spirosperma. 



EMBOLOCEPHALUS YAMAGUCHIl (BRINKHURST, 1971) (CLITELLATA: TUB1F1C1DAEI 

FROM SOUTH AUSTRALIAN STREAMS 



AM P1NDER & PK McEVOY 



PIN'DER. AiVl & McEVOY, PK. 2002 Embolocephalus samaguchii (Brinkhursi. 1971) 
(Cluellata: Tubificidae) from South Australian streams. Records of the South Australian 
Museum S$(2): 139-145. 

Papillate tubificids from sucams in the Mount Lofty Ranges are identified as 
hmholnrepludus ynmoguchii (Brinkhurst, 1971) and represent the first records of this largely 
Holarctic genus from Australia. Collection details arc provided for records of a second papillate 
species, identified as belonging to the genus Spirosperma 

AM Pindcr. Department of Conservation and Land Management, PO Box SI, Wanneroo, 
Western Australia 6946. PK MeEvoy, Australian Water Quality Centre. Private Mail Bag J, 
Salisbury, South Australia 5J08. Manuscript received October 2001. 



Must tubificids with papillate body walls are 
contained within a complex of genera 
(Including Embolocephalus Randolph and 
Spirosperma Eisen) established, or re- 
established, following revision of the genus 
Peloscolex Leidy (Brinkhurst 1991; Brinkhurst 
& Wetzel 1984; Holmquist 1978, 1979). These 
eenera h3ve a largely Holarctic distribution, 
with few records from southern continents. The 
latter consist of a record of the otherwise 
European Embolocephalus veluitnus (Grube, 
1879) from Venezuela (Dumnicka, 1983), an 
identification questioned by Brinkhurst and 
Marchcse (1989), and papillate tubificids of 
uncertain idcnlity from Peru (Brinkhurst & 
Marchese 1989). The North American species 
Quistadrilus multisetosus has been recorded as 
far south as Mexico. In Australia, Timms ( 1 97S j 
recorded Peloscoiex (sic) from a lake in 
Tasmania and Pindcr & Brinkhurst (1994, 2000) 
noted papillate tubificids from Victoria and 
New South Wales. The latter were all immature 
so descriptions were not possible, but they were 
tentatively assigned to Spirosperma, based on 
the presence of papillae and the form of the 
chaetac. Collection details for this species, 
including the first records from South Australia, 
are presented below for the first time. Other 
specimens, collected from streams in the Mount 
Lofty Ranges in .South Australia, are herein 
described and identified as Embolocephalus 
yamaguchii (Brinkhurst, 1971), a species 
otherwise known only from Lake Btwa in 
Japan. 



Methods 

Serially sectioned specimens were cut at 6 uni 
and stained in haematoxylin and eosin. 
Measurements were taken using an eyepiece 
graticule on a Zeiss Jenamed 2 compound 
microscope calibrated with an Olympus stage 
micrometer. Abbreviations used in the line- 
drawings are a: atrium, cp: cuticular pad, e: large 
epidermal papillae, ed; ejaculatory duct, ff: female 
funnel, mf: male funnel, p; prostate, pe: penis, sa: 
spermathecal ampulla, sc: spermathecal chacu, 
scg spermathecal chaetal gland, v: vas deferens. 
Specimens are either in the senior author's 
collection (AP) or Ihe Australian Water Quality 
Centre collection (AWQC); returned to WSL 
Consultants (WSL) or Australian Water 
Technologies (AWT); or deposited with the South 
Australian Museum (SAM) or the Museum of 
Victoria (NMV). 



Systematics 

Embolocephalus Randolph, 1 892 

Type species 
Embolocephalus velutinus (Grube, 1879) 

Diagnosis 

From Holmquist (1978) and Brinkhurst (1981). 
Prostomium and sometimes first segments 
retractable within the rest of body. Body wall with 
epidermal papillae, generally concealed by a 



140 



AM PINDER & PK McEVOY 



secreted layer with embedded foreign particles. 
Hair chaetae present dorsally, usually broad and 
sabre-like, with pectinate crotchet chaetae. Bifid 
and/or simple pointed chaetae present ventrally, 
usually both in anterior bundles. Spermathecal 
chaetae modified on X, lying in a well-developed 
chaetal sac. Male ducts each with a long winding 
vas deferens, usually narrower entally and broader 
(up to twice the width) ectally, entering the atria 
apically. Atria long and tubular to crescentic, 
broadest near the prostate union, tapering to a 
short ejaculatory duct. One large prostate gland 
joining each atrium towards the middle of the 
latter. Penes well developed without thickened 
cuticular sheaths. Spermathecae with ovoid to 
oblong ampullae, well set off from the ducts, with 
pores usually more or less in front of and lateral 
to the spermathecal chaetal sacs. 

Distribution 

Europe, North America, Japan, Kamchatka, 
?VenezueIa and now Australia. 



Embolocephalus yamaguchii (Brinkhurst, 
1971) 

Peloscolex yamaguchii Brinkhurst, 1971: 505 
Peloscolex sp. Yamaguchi, 1953: 295 
Embolocephalus yamaguchii (Brinkhurst, 1971) 



Ohtaka, 1994: 52; 1995: 174 



Material examined 

South Australian specimens. SAM E3095- 
3101: 3 mature specimens serially sectioned, 3 
mature dissected and 1 immature whole-mounted, 
Aldgate Creek at Strathalbyn Road, Mylor, 
tributary of the Onkaparinga River (AWQC site 
3217), 35°02'S 138°45'E, samples E33, 3 June 
1997 and F32, 27 Oct 1997. AP: 1 mature serially 
sectioned, collected as above. SAM E3 102-3 103: 
1 mature in alcohol, Onkaparinga River at 
Houlgraves Weir (AWQC site 3205), 35°05'S 
I38°43'E, sample E37, 3 June 1997, and 1 mature 
whole-mounted from same site, sample F37, 
28 Oct 1997. SAM E3104: 1 mature in alcohol, 
Aldgate Creek at Aldgate Valley Road (AWQC 
site 13022), 35°02'S 138°45'E, sample H41, 2 
Dec 1998. AWQC: 5 immature in alcohol, 
Onkaparinga River at Hack Bridge (AWQC site 
3212), 35°03'S 138°45'E, sample H43, 1 Dec 
1998. AWQC: 1 immature in alcohol, Lenswood 
Creek at gauging station (AWQC site 3208), 



34°56'S 138°50'E, sample OCB148, 17 Jul 2000: 
AWQC: 2 mature in alcohol, Onkaparinga River 
at Silver Lake Road (AWQC site 13023), 35°04'S 
138°45'E, sample OCB181, 17 Oct 2000. 
Collections by V. Tsymbal, P. McEvoy, A. Lang 
and S. Wade (Australian Water Quality Centre, 
South Australia). 

Japanese specimens. From collection of A. 
Ohtaka (Hirosaki University, Japan): 1 mature 
specimen serially sectioned, 1 mature whole- 
mounted and 2 mature in alcohol (now dissected), 
Lake Biwa, Japan. 

Habitat 

This species has been collected only from a 
sixth order reach of the Onkaparinga River 
upstream of Mount Bold Reservoir and from two 
fourth order tributaries, between 245 and 295 
metres above sea level. Specimens were collected 
from both riffle and edge habitats, with substrates 
ranging from those composed mainly of bedrock, 
boulder or cobble to those dominated by finer 
sediments and/or detritus. Willow root mats are a 
common feature on the stream bed of most sites. 
Riparian zones had an overstorey of either 
indigenous Eucalyptus species and Acacia 
melanoxylon and/or introduced willows (Salix 
spp.). Land uses include grazing, horticulture and 
urban settlement. Apparently similar habitats in 
adjacent catchments were sampled with the same 
sampling intensity without finding E. yamaguchii. 

Description of Australian specimens 

Length of preserved specimens 15.6-25.5 mm, 
width at clitellum 0.7-1.1 mm. Each pre-clitellar 
segment with a band of tall, broad epidermal cells 
forming a raised transverse ridge in the middle of 
the segment, pronounced dorsally and laterally but 
virtually absent ventrally. Body surface with 
foreign material adhered to elongate ovoid 
papillae covering the entire body except for the 
clitellum (Fig. 1). Papillae mostly 15-22 urn long 
on pre-clitellar segments (measured on the longest 
axis, including foreign particles) and 20-34 urn 
long on post-clitellar segments, and mostly 17- 
25 urn high (with most of the height constituted 
by the foreign material). Ridges of anterior 
segments with fewer or no papillae. Each segment 
also with 2 rings of papillae that are taller and 
broader than normal (up to 40 um wide and 
40 um high including foreign material), which 
give the appearance of darker (or sometimes 
lighter) bands around each segment, 1 ring about 
at the level of the chaetae and 1 anteriorly on each 
segment, most pronounced on posterior segments. 



IMnoLOCEPIIALVS YAMAGUCH11 KROM SOUTH AUSTRALIAN STREAMS 



141 




FIGURE t. /:'. ytinuiguchii A. Enlarged view of body 
wait of BegfttySttlS towaids the posterior end of SAM 
>i3lM8 (prior to slide mounting) showing normally sized 
and enlarged (e) papillae; B, Portion of SAM E3I04 
showing lateral view of arnerioi segments (lop), 
clilellum (centre) and post-chiellar segments (bottom). 
Scale bar for A 100 um. B 500 urn 



Other (possibly sensory) papillae present with the 
foreign materia! adhered only to the sides, with 
the tap of the papillae exposed, arranged in ill - 
defined rings around each segment (about 10 of 
these papillae per ring), usually 1 ring at the level 
of the chaetae and I or more rings between die 
ehaetae and intersegmental furrow. These partially 
naked papillae are about Hie same sue as Ihe 



shorter type of fully encased papillae and are 
.sometimes visible as darker or paler spots. A pad 
of tall, thin epidermal cells, with cuticle 10 times 
thicker (up to 17 urn) than elsewhere, present 
medially on the spcrmalhcca! segment, adjacent to 
the anterior spermathecal chaeta. Clilellum 
covering posterior half of X arid all of XI, paler 
ihan other segments due 10 a smooth even coaling 
of soft while granular material (Pig. 1 ), 

Prostomtum partially retracted into first 
segment in fined animals. Pharynx with dorsal 
muscular pad in Ii to HI, with pharyngeal gland 
ceils on pharynx and on septa 3/4 to 4/5. 
Oesophagus from IV to VI, widening mio broader 
Ihickcr-wallcd intestine in VII. 

Ventral chaetae 130-211 urn long and 7-14 urn 
wide at nodulus, generally smallest in pre-clitellar 
bundles. Ventral bundles on 11 to VIII usually with 
I bifid and I simple pointed chaeta, rarely I extra 
bifid chaeta and/or 1 or 2 extra simple pointed 
chaetae. Bifid chaetae with teeth equal in width 
but upper tooth up to 1.5 times longer than lower 
(Fi.g. 2B). Posterior ventral bundles with 1 (rarely 
2) bifid chaetae, with upper teeth becoming 
shorter than lower and lower tooth greatly 
expanding in width and becoming notably 
recurved (Fig. 2C). Dorsal chaetal bundles 
normally with 2 to 4 slightly curved hair chaetae, 
220-420 pm long and 4—7 urn wide, with finely 
plumose shafts and blunt, slightly swollen, tips, 
with an equivalent number of short (32-75 urn 
long and 1-2 pm wide) finely pectinate crotchets 
with thin parallel outer teeth (Figs 2D.E). 
Posterior dorsal bundles with fewer of both types 
of ehaetae. Chaetae of spermathecal segment 
modified (Figs 2A, 3A,B), ventral ehaetae absent 
in XI. 

Genitalia paired (Figs 2, 3B). Testes 
anteroventral in X, ovaries anteroventral in XI 
Male funnels large, ciliated vasa deferentia broad 
near funnels (35 pm), narrowing shortly thereafter 
(17-20 pm) and widening slightly ectally 
(25-33 pm), highly coiled, joining ental end of 
atria. Atria each with a narrow lumen and layer of 
tall glandular lining cells, ercsccntic, broadest 
where a. single prostate gland joins (slightly entad 
of medial), tapering at either end, the ectal end 
forming a short ejaculatory duct which enters the 
penis apically. Atrial muscle layer thin, wider 
where proslale joins. Atrium, including 
ejaculatory duct, measuring 800 x 1 10 pm to 1000 
x 185 pm. Prostate glands voluminous, sometimes 
ballooning into following segment. Penes broad 
entally. narrowing at about one-third of the 
distance from the tip to form a narrower ectal end. 



142 



AM PINDER & PK McEVOY 




FIGURE 2. E. yamaguchii. A, Reconstruction of genitalia (with organs drawn in the same plane for simplicity) 
from serial sections of SAM E3095; B, tips of anterior ventral chaetae; C, tip of posterior ventral chaeta; D, dorsal 
pectinate chaeta; E, tips of dorsal hair chaetae, with plumosity shown for one hair. Scale bar for A 200 urn. 



penes within thin-walled muscular penis sacs 
attached to the dorsolateral body wall by 
numerous muscle fibres. Penes and penis sacs 
with cuticle no thicker than on body wall (ie 
without penis sheaths). Male pores in line with 
ventral chaetae on anterior half of XI, within a 
common transverse depression of the ventral body 
wall. Spermathecal ampullae variable in size 
(770-1350 urn long), each containing 1 or 2 long 
thin spermatozeugmata (length of only one whole 



spermatozeugmata measured, 660 x 45 urn, but 
others 80 um wide in cross-section, indicating 
greater length). Spermathecae with stout ducts 
(300-370 um) that are constricted ectally, leading 
to pores anterior on X slightly medial to line of 
ventral chaetae. Spermathecal chaetae long (510- 
670 um) and thin (10 um) with grooved tips, ectal 
half lying in well-developed chaetal sacs. The sacs 
tall (220-300 um) and muscular, with inner lining 
tissue that can be everted with the tip of the chaeta 



EMBOLOCEPHALUS YAMAGUCHII FROM SOUTH AUSTRALIAN STREAMS 



143 




VIII IX 



FIGURE 3. E. yamaguchii. Reconstruction of genitalia (with organs drawn in the same plane for simplicity) from 
serial sections of A, SAM E3096 and B, SAM E3097. Scale bar for A 200 urn. 



to form a large papilla and with muscular layer 
extending to cover the ental end of the chaeta. Sacs 
with a gland attached dorsally, varying in extent of 
development (between and within individuals). 
Spermathecal chaetal sacs located asymmetrically, 



both opening slightly ventral to the line of somatic 
ventral chaetae, I immediately posterior to 
spermathecal pore on one side and the other close 
to 1 1/12 in front of the sperm funnel on other side. 
Female funnels posterolateral on XI. 



144 



AM PINDER .V: PK McFVOY 



Remarks 

The presence of simple-pointed chaetae on 
segments 11 to Vlll, separate openings for the 
spcrmalnceal duels arid sperinalheeal ehaelae, 
dorsolateral ridges on anterior segments and 
asymmetrical location of the spenmathccal chaetae 
clearly ally these Australian specimens with the 
Japanese species £. yamaguchii rather than with 
any of its congeners We initially considered that 
the Australian specimens were either conspecific 
with E, yamaguchii or represented a very similar 
sisler-species. The. latter was seen as more likely 
since E. yamaguchii is known only from Lake 
Biwa in Japan and other Embolocephalus seem to 
have fairly restricted Holarciic distributions 
However, a thorough comparison of E. 
yamaguchii (from the descripiion in Ohtaka 1994 
and material seen by us) and the Australian 
specimens led to the conclusion that there was no 
morphological justification for a new species 
description. The only differences between the 
Australian and Lake Biwa specimens are the size 
of the worms and the size of some genital organs. 
None of the Australian specimens reach rhe 
maximum length of 35 mm measured for the £. 
yamaguchii neotypes (Ohtaka 1994), but most 
Lake Biwa specimens are less than 30 mm 
(Ohtaka pers. coram.) The vasa deferenlia ol Ihe 
Australian specimens are narrower than those (20- 
24 um wide entally, 40 um ectally) recorded for 
E, yamaguchii by Ohtaka (1994) and the ectal 
portion is not so enlarged relative to the ental 
portion. However, the vasa deferentia of Lake 
Biwa specimens measured by us (as narrow as 
13 pm entally to a maximum of 25 um ectally) 
more closely match measurements from the South 
Australian material. Other components of the 
genitalia (atria, spermathecal ampullae and 
spermathecal chaetae) are slightly larger in the 
Australian specimens than recorded by Ohtaka 
(1994.). The raised epidermal pad was not noted 
for E. yamaguchii by Ohtaka (1994) but was 
present on Lake Biwa material seen by us and is 
visible in Fig. 4B of Ohtaka (1994i. This feature 
is not known for other Embolocepholus species. 

These specimens represent the first records of 
this species and genus in Australia and one of 
the few records of the genus from southern 
continents. The known distribution of this 
species (Lake Biwa in Japan and the Mount 
Lofty Ranges in South Australia) is exceptional 
within the Australian tubificid fauna. Other 
tubificids found in Australia are either endemic 
or cosmopolitan (occurring in most, if not all, 
other continents). 



'ISpirasperma sp. 

Material examined 

All specimens immature, WSL consult 
voucher AN30: 2 in alcohol. Old Namoi River at 
Bullerawa Station, (Duncan's Junction) New 
South Wales. 3<m'()5"S l49 o 04'00 n E 27 Jun 
2000 AWT: I specimen, Georges River ;>t 
Cambridge Avenue. New South Wales, 33°58'S 
I50°54 , E. I Apr 1996: I specimen. Peach Tree 
Creek, at Weir Reserve, New South Wales, 
33°45"S 150°4rE. 17 Jul 1996; 1 specimen. 
Second Ponds Creek downstream of Rouse Hill 
Sewage Treatment Ponds, 33 o 40'S l50 o 55'E. 
1 Apr 1996. AP: I whole-mounted on slide. 
Plenty River at Lower Plenty. Victoria, 37*44 'S 
14.VWE, 10 Apr 1994, coll S. Schreibcr 
(formerly Monasli University); Glenelg River at 
RocMands, Victoria, n°l4'S !4r'57'E. 3 Jun 
1994, coll. S. Schreibcr. NMV F8186S: I in 
alcohol, LaTrobe River at Mot-Willow Grove- 
Road Bridge, Victoria, 3S°1 1' 18 "S 146"L512 h. 
NMV F81864: 1 in alcohol, Curdies River. 
Victoria. 38"20'S I43"08'E. 7 Feb 1992, coll G. 
Quinn (formerly Monash University). AWQC 
voucher 5665; 1 in alcohol. Pilbv Creek outlet 
(AWQC site 13095), South Australia, 33"59 , S 
140°53E, 24 Nov 1998. coll. D. Schulze and T. 
Venus. 

Brief descripiion 

Prostornium retractable within mouth. Bndy 
wall densely papillate, with foreign material 




to 



FIGURE 4 Spirospcrma sp A, ventral chaetae of 
anterior to mid-body segments; B, ventral chaela of 
pOdWtiOr segment; C. tips of dorsal crotchet chaetae. 



EMBOLOCEPHALUS YAMAGVCHII FROM SOUTH AUSTRALIAN STREAMS 



US 



attached to most papillae. Some papillae with 
foreign matter on lateral walls only. Without rings 
of distinctly larger papillae. Ridges of enlarged 
epidermal cells absent on anterior segments. 
Ventral chactae (Fig. 4) all bifid, 3 to 6 pcT 
bundle, with upper teeth 2 lo 3 limes as long as 
lower teeth in anterior and mid-body segments, 
1.5 times as long as lower teeth in posterior 
segments (Fig. 4). Ventral chaetae of first few 
segments usually slightly longer and thicker than 
the rest. Dorsal bundles with 1 to 3 smooth hair 
chaetae and I lo 3 very small crotchet chaetae 
with finely bifid ends and 1 or 2 intermediate 
teeth (Fig. 4). 

Remarks 

This species lacks simple-pointed chaetae 
anteriorly and so has been tentatively identified as 
a Spirosperma rather than an Emhnlocepluilus, but 
mature specimens arc required to confirm the 
generic affiliation, ft appears to be widespread in 
rivers of southeastern Australia. 



Acknowledgments 

Specimens of £ yamqguchli were collected as pan 
of the Monitoring of Rivet Health Initiative funded by 
the Land and Water Resources Research and 
Development Council and Environment Australia. 
Funding from the Wildlife Conservation Fund of the 
Department for Environment and Heritage (South 
Australia) enabled identification to species of 
oligochaetes from the above project. Laboratory and 
microscope facilities were made available by Stuart 
Halse (Department of Conservation and Land 
Management). Specimens of Spirosperma were made 
available by Australian Water Technologies, Alena 
Olaistet at Monash University and Kylie Swinglcr at 
WSL Consultants (specimens from a Department of 
Land and Water Conservation NSW project). Akifumi 
Ohtaka (University of Hirosaki, Japan) kindly 
provided material of E. yamaguchii from Lake Biwa 
and engaged in useful discussions with the authors. 
Gordon Thomson (Murdoch University, Perth) 
performed the serial sectioning and Jane McRac 
(Department of Conservation and Land Management) 
assisted with photography. 



References 



Brinkhurst, RO 19V I. A phylogenetic analysis of the 
Tubificinae (Oligochacta: Tubificidae). Canadian 
Journal of Zoology 69: 392-397. 

Brinkhurst, RO. 1981. A contribution to the taxonomy 
of the Tubificinae (Oligoehaeta: Tubificidae) 
Proceedings of the Biological Sociers 1 of Washington 
94: 1048-1067. 

Brinkhurst, RO & Marchese, M 1989. 'Guide to die 
freshwater aquatic Oligochacta of South and Central 
America'. Asoclaeion Ciencias Naturclcs del Litoral: 
Santo Tome, Argentina 

Brinkhurst, RO & Wetzel. MJ. 1984. Aquatic 
Oligochaeia of the world Supplement. A catalogue 
of new freshwater species, descriptions and revisions. 
Canadian Technical Report of Hydrography and 
Ocean Sciences 44: I -10 1. 

Dumnicka, E. 1983. Tubificidae (Oligoehaeta) from 
subterranean waters, with description of two new 
genera. Bijdragen tot de Dierkundc 53: 255-261 

Holmquist, C. 1978. Revision of the genus Peloscolex 
(Oligoehaeta: Tubificidae) I. Morphological and 



anatomical scrutiny; with discussion on the genetic 
level. Zoologica Scriptal: 187-208. 

Holmquist. C. 1979. Revision of the genus Peloscolex 
(Oligoehaeta: Tuhificidae) 2. Scrutiny of the species. 
Ztiolngwa Scripia 8' 37-60. 

Ohtaka. A. 1994. Redescription of Emlwlocephalus 
yamaguchii (Brinkhurst, 1971) comb. nov. 
(Oligochaeia: Tubificidae). Proceedings of the 
Japanese Society of Systematic Ziudogy 51: 34-42. 

Pinder. AM & Brinkhurst, RO. 1994. 'A Preliminary 
Guide to the Identification of the Microdnle 
Oligoehaeta of Australian Inland Waters'. 
Cooperative Research Centre for Freshwater 
Ecology; Albury. 

Pinder. AM & Brinkhurst. RO. 2000. A review of the 
Tuhificidae (Annelida: Oligochaeia) from Australian 
inland waters. Memoirs of Museum Victoria 58: 39- 
75. 

Timms, BV 1978 The benthos of seven lakes in 
Tasmania Archiv fiir Hvdrohiologiv 81: 422-444 



EARLY ABORIGINAL FISHING TECHNOLOGY IN THE LOWER 

MURRAY, SOUTH AUSTRALIA 



Philip A. Clarke 



Summary 

The early Aboriginal fishing technology of the Lower Murray region of South Australia is described 
and compared with other areas in south-eastern Australia where fish was a dominant dietary 
component of hunters and gatherers. This is a study of cultural geography, the chief concern being a 
description of hunting and gathering techniques and their significance to Aboriginal occupation of 
the landscape. 



EARLY ABORIGINAL FISHING TECHNOLOGY IN THE LOWER MURRAY, 

SOUTH AUSTRALIA 



PHILIP A CLARKE 



CLARKE, PA. 2002. Early Aboriginal fishing technology in the Lower Murray, South 
Australia. Records of the South Australhm Museum 35(2): 147-167. 

The early Aboriginal fishing technology of" the Lower Murray region of South Australia is 
described and compared with other areas in south-eastern Australia where fish was a dominant 
dietary component of hunters and gatherers. This is a study of cultural geography, the chief 
concern being a description of hunting and gathering techniques and their significance to 
Aboriginal occupation of the landscape. 

PA Clarke. Science Division, South Australian Museum. North Terrace, Adelaide, Soulh 
Australia 5000. Manuscript received 7 December 2001. 



Introduction 

Aboriginal fishing technology in the Lower 
Munray region is discussed from the pre-European 
period io the early years of European settlement. 
A major aim is to describe how hunting and 
gathering techniques used by Lower Murray 
people made their region culturally distinctive 
(Fig. I). 1 This region is defined as the area from 
Cape Jervis in the west, east to Wellington on !he 
Murray River and south to Kingston, taking in (he 
whole of the Lower Lakes, Coorong and 
associated coastal belt It is estimated that just 
prior to European settlement, the region supported 
a population of 5000 Aboriginal people, although 
this figure was probably seriously affected by two 
early waves of smallpox (Brown 191S: 230; 
Campbell 2002: 119-133; Clarke 1994: 57-63; 
1995: 156, footnote 1; Gale 1969). The 
descendants of these people, many of whom still 
live in the region, generally call themselves 
Ngarnndjeri (Bcrndt & Bemdt 1993; Clarke 1994; 
Hemming & Jones 2000; Jenkin 1979). ; 

In the Lower Murray region ihe Murray River 
Hows from Murray Bridge to Lake Alcxandnna 



through an open valley cut across a very low and 
flat limestone karst plain, which is less than 30 
metres above sea level (Fenner 1931: 81-83: 
Twidale 1968. 148-149. 383-384). Below 
Wellington, the river becomes two large Jakes 
(Alexandrina and Albert) and a series of channels 
in the form of a delta, eventually exiting behind 
scattered islands at the Murray Mouth ' Here, the 
river meets the Coorong, which drained the South 
East region of South Australia before European 
intervention. 4 The sea, winds and tides combine to 
drive the river back with heavy sand dune 
systems, called Sir Richard and Younghusband 
Peninsulas (northwest and southeast parts, 
respectively).' From the point of view of the early 
Aboriginal inhabitants, the delta of the Lower 
Murray provided the region with many kilometres 
of shoreline for hunting and gathering activities. 

The climate of the Lower Murray region is 
influenced by the powerful 'controls' of its 
temperate latitude, the proximity Of the sea and 
the relief of the land (Fenner 1931: 125; Howchin 
1909: 142, Penney J983: 85-93; Schwerdtfeger 
1976: 75-86). Upstream, long sections of both the 
Murray River and its main tributary, the Darling 



Hunting and galhetmg practices in the region since ihe I ( J40s are discussed elsewhere iCIajVe 21X12). 

I he ISganitidjcri ( - Nartinycri) were formerly made up of deseem groups who spoke one of several dialects, such as Ramindjcri Yaraldi 

t. = Jarildekald) and Tungiuti { = Tanganekald). 

Lake Alexandrina is called 'Lake Victoria* on some earlynffici.il maps (Cockburn 1984:7). 

The Coorong was formerly known as ihe 'Souu\ Fasi Brunch' in nfejencc (u Lake Alc.sandrina. which it joined ill Pelican Poiui (Cuiquc |R. 

Penneyl in the Soulh Australian Mu%atirte. September IfM2, vol 2; IK-231 

Otic to Ihe action of (he ocean currants, (lie lr«:siinn of ihe i ivei esil into the sea is constandy moving; ut present il is migrating norUiwards towards 

Goolwa at the raw of several metres per year <P. Tuckwell, peri coinm.l. The complete disappearance of Barker Knoll at ihe Mouth as early as 

1 85° 'Linn 1988, 78 i indicates dial some movement is a natural feature. 



148 



PA CLARKE 



^URRAY BRIDGE 

TAILEM BEND 




J 

KINGSTON 



FIGURE 1 . The Lower Murray cultural region. 



EARLY ABORIGINAL HSHINC. TBC1INOUXJY 



t« 



River, flow through semi-arid regions. 
Nevertheless, the Lower Murray region is entirely 
contained within the high rainfall area of South 
Australia, receiving 350-750 mm per year. [I 
comes under the rainfall shadow of the Mount 
Lofty Ranges to the west, with precipitation also 
increasing near the coast- Annual average 
temperature, throughout the region is less than 
I8°C, with the greatest range of temperatures 
heing during the summer months (Fenner 193] 
65, 126: Griffin & McCaskill 1986; 50-51, Laut 
ctal 1977) 



Stumers of Aboriginal Environment Use Dat\ 

The literature of Aboriginal hunting and 
gathering technology for the region is based on 
Tour main sources of data — the archaeological 
record, the histoiical ethnographic record, 
scientific analysis of the properties of naturally 
occurring substances and contemporary research 
with Aboriginal people. This paper deals mainly 
with historical ethnographic sources of 
information, which includes artefacts collected 
from living people. Archaeological investigations 
provide evidence of Ihe importance of fish in the 
diet of coastal/riparian Aboriginal groups, with an 
indication of the material cultural items and the 
main species fished. Biological analysis of htiman 
bone also assists in determining the pre- Burnt e W 
diet (Pate 1997 1998, 2000). Scientific methods 
of analysis ol food sources, such as those 
determining the. pharmacological and nutritional 
properties, can illustrate their putential human 
uses. The literature investigating the usefulness of 
Australian fish as food and medicine includes 
Brand Miller el al (1993: 222-223), Clarke (1989: 
3) and isaacs (1987: 153-164). Nevertheless, 
cultural perceptions influence the potential use of 
plants and animals, wilh nol all availahle 
resources being fully utilised. To understand how 
Lower Murray Aboriginal people used their 
environment, and thereby moulded their cultural 
landscape, we must at present rely primarily on 
records made by early European observers, 
supplementing this with ethnographic data from 
contiguous nparian/coasial regions. 

Although useful data on pre- European plant 
use, such as species identification, method of use 
and seasonality, can be obtained from 



contemporary ficldwork with Aboriginal people, 
there are significant post-European changes in 
Lower Murray Aboriginal culture and in tin- 
physical landscape itself. For instance, 
information received from Lower Murray 
Aboriginal people in recent times on bush foods 
used during the last 50 to 60 years indicates Tar 
less use of roots than before (Clarke 1988: 64) In 
contrast, other indigenous foods such as fish, 
waterfowl, kangaroos, emus and berries continued 
10 be used in restricted contexts. Nevertheless, the 
introduction of foreign species, such as European 
carp, and environmental changes in the waterways 
have decreased numbers of some Australian fish 
species. The availability of European- type foods, 
especially flour, potatoes and canned foods, 
obtained from missions, farm stations and towns 
led to a decline in many indigenous food sources. 
The decrease in 'bush food' consumption was 
particularly marked for those species requiring 
significant labour expenditure Other foods were 
replaced by European varieties with more 
favourable properties. The bulrush root, for 
example, contains a great deal of fibre that makes 
eating difficult. Unlike earlier Aboriginal foods, 
many European sources were available at all limes 
of the year because they were imported 

The detailed recording of Aboriginal culture in 
the Lower Murray region started with the German 
Heinnch A- E. Meyer, who ran a mission at 
Encounter Bay during the late 1830s and early 
1840s (Meyer 1843, 1846). From him we obtain a 
Ramindjeri-speaker's view of Lower Murray 
culture. The Englishman George Taplin 
established an Aboriginal mission at Point 
McLeay on the south-western shore of Lake 
Aiexandrina in 1859. fn the 1870s he published 
two main books on Ngarrindjcri ( = Namnycri) 
culture, based on records in Ws journals (Taplin 
1S59-79, 1874, 1879). Taplin used Rammdjeu 
sources as a guide to his research, although local 
Aboriginal groups living around Point McLeay 
were mainly Yataldi-speakers. He described 
people who were feeling the impact of European 
expansion and the considerable changes it had 
brought upon them and the Lower Murray 
landscape, particularly at the northern end. One ul 
Taplin's initiatives to improve their circumstances 
was to establish a local fishing industry for 
Aboriginal people at the mission (Jenkin 1979: 
97-98. 110-1 ll). 7 



I or an archaeological overview ol Hie Lower Murray / Sown Hasi tSisfffcis. sec Campbell l|9U. 1 r '39. IWi. t'aiiiphcll el «) il'-iiAi Lusbtaft 
O u 78|atKll*rciil eislO'. I) 

Trim arc many v-rttries in Taptin's journals concerning Hie establishment, of UV Bs&fitf industry Tire MpAitUII irMeiriice*; an; 71 NefaemS/f IKVJ. 
l&Otlohd iKSQ.2i November 1859. 21 December J&59. II hman I Hfa 7 Fehi uary I3SO. 15 Irbrunry IMMUwl I March Ifcoll 



150 



PA CLARKE 



Adding to the missionary records in the 19th 
century are the recordings of colonists such as 
George French Angas (1847a, 1847b), William A. 
Cawthorne (1844 [1926]), Dirk M. Hahn (1838- 
39), Richard Penney (1840-43) 8 and William 
Wells (1852-55). During this century, various 
anthropologists studied Lower Murray Aboriginal 
culture: Ronald M. Berndt (1940; with Catherine 
H. Berndt 1993), Alison Harvey (1939ms, 1943) 
and Norman B. Tindale (1930-52, 1931-34, 
1934-37, 1935, 1937, 1938, 1938-56, 1941, 
1981, 1987; with Mountford 1936; with Pretty 
1980). Their accounts are rich in detail concerning 
past hunting and gathering practices, filling many 
gaps that were left in the early historical record. 
They also contain examples of post-European 
innovations by indigenous people. Largely due to 
the interests and efforts of the Director of the 
South Australian Museum, Edward C. Stirling, 
and Ethnology Curator, Norman B. Tindale, the 
South Australian Museum possesses a large 
collection of early artefacts from the Lower 
Murray region. 9 In the 1980s the present author 
commenced studying Aboriginal relationships to 
the physical landscape (Clarke 1985a, 1985b, 
1986a, 1986b, 1987, 1988, 1989, 1994 chapter 4, 
1998, 1999b, 2001a). The region and Aboriginal 
relationship to it also featured in the Hindmarsh 
Island Bridge controversy (Bell 1998; Stevens 
1995; Wilson 1998). The attention that scholars 
have directed towards the Lower Murray has 
meant that it is ethnographically one of the best- 
described regions in southern Australia. The 
biases within this literature are discussed 
elsewhere (Clarke 1994 section 11.3, 1999a: 149- 
151,2001a: 19-20). 



Early Aboriginal Fishing Technology 

In the pre-European period the high Aboriginal 
population along the river frontage and the coastal 
zone of South Australia was in sharp contrast to 
the sparse population in the interior. In the Murray 
Basin region the distribution of people was 
estimated to be 0.4-0.5 square kilometres for each 



individual, in contrast to 31-88 square kilometres 
required in Central Australia (Jones 1974: 326; 
Lawrence 1968: 72-73; Maddock 1972 [cited 
Rose 1987: 22-23]; Meggitt 1962 [cited Abbie 
1976: 46-47]; Smith 1980: 68-90; Tindale 1981: 
1860). It has been demonstrated that the physical 
environment of the Lower Murray formed both a 
natural and cultural region (Clarke 1994). Stable 
isotope research on human bone indicates that, in 
general, neither people nor foods were moving 
from the Murray Bridge area to the adjacent 
Murray Mouth and Coorong (Pate 1998). The 
Lower Murray therefore formed a separate hunting 
and gathering region. Nevertheless, some aspects 
of the material culture and natural resource use by 
Aboriginal people here was similar to that of 
riverine and coastal communities described from 
elsewhere in Australia (Hallam 1975; Lawrence 
1968: 85-122; Lourandos 1997: 195-243). 

In the 1930s Tindale mapped many of the 
favourite fishing places and lookouts in southern 
South Australia, from informants such as Louisa 
Eglinton (Narangga woman), Milerum (Clarence 
Long, Tangani man) and Albert Karloan (Yaraldi 
man)."' In the Tangani language, elevated areas 
used as 'watch places' were called popatdi, 
whereas in Ramindjeri they were koinkoinj." 
Older men generally used these when looking out 
for fish shoals and bird movements, as well as for 
monitoring the movements of their neighbours by 
observing the smoke from their campfires. In 
comparison to adjacent regions, such as the 
Murray Malice and Mount Lofty Ranges, the 
Lower Murray landscape offered Aboriginal 
people abundant opportunity to make fish a more 
significant food source. 

Before the arrival of Europeans, Aboriginal 
people in southern Australia did not widely use 
the fishhook and line (Curr 1883: 110; Davies 
1881: 129; Eyre 1845, vol. 2: 266-267; Massola 
1956; Meyer 1846 [1879: 192]; Olsen 1991: 5). 12 
Nevertheless, the use of bone bi-points or fish 
gorges (muduk) and fishing lines has been 
recorded from along the Murray River (Flood 
1999: 48; Gerritsen 2001; Pretty 1977: 321-322). 
Similar items have been recorded from the 



Newspaper articles by Richard Penney between 1840 and 1843 are reprinted in the Journal of the Anthropological Soc/ery of South Australia 1991- 
1-107. 

From 1988 to 1999, a major exhibition on Aboriginal culture in the Lower Murray, called 'Ngurundcri, a Ngarrindjeri Dreaming', was open at the 
S.A. Museum (Hemming & Jones 2000). 

Tindale Aboriginal Site Maps of southern South Australia, South Australian Museum Archives. Adelaide. Also see Tindale (1935-60: 15). 
Tindale (no date, 'Milerum', Stage A, #9, Archives, South Australian Museum, Adelaide). 

Tindale (1934-37: 285) recorded a description by Karloan (Karlowan) of fishhooks being made from bone, gum and stick, but this was possibly a 
post-European development. 



EARLY ABORIGINAL FISHING TECHNOLOGY 



151 



Gippsland and Geelong areas of southern Victoria 
(Smyth 1878 vol. 1: 391). Shell fishhooks have 
also been recovered from coastal shell middens in 
eastern Victoria (Mulvaney & Kamminga 1999; 
292). 

After Europeans arrived, Aboriginal people in 
the Murray River and Lower Murray regions 
adopted new fishing techniques and appeared to 
have made their own version of the fishhook." In 
1938 Tindale recorded Albert Karloan who said: 

Our fish hook was made from a bone of a 
kangaroo tied like a real fish hook to another 
piece which was the point but our old folks used 
ihe fish spear all (lie time: nothing was as good 
.is Ihe real fishing spear! V/e walked along the 
banks & got fish in the reeds (Tindale 1930-52: 
42). 

Kuropean twine, hooks and lines were also 
handed out at the Point McLeay Mission (Taplin 
Journals: 18 September 1862). Fishing techniques 
used in the Lower Murray region included netting, 
spear and club fishing, trapping and opportunistic 
collccting. 



Net-Fishing 

The Ngarrindjeri people referred to the aci o| 
ne.timg fish as ngerin (Taplin 1879: j 30). Marine 
net fishing often involved the coordination of a 
large number of people Worsnop provided a vivid 
account of Aboriginal people sea fishing in Ihe 
19th century. He records. 

In Encounter Bay I have seen the natives fishing 
almost daily Two parties of them, each provided 
with a large net, square in form, with a stick at 
either end, and roiled up, swam out a certain 
distance from the shore, and then spread 
themselves out into a semicircle. Every man 
would then give one of the slicks round which 
his piece of not was rolled to his right hand man. 
receiving another from his left hand neighbour, 
bunging the two nets together, thus making a 
great seine. They now swam in towards the 
shore, followed by others of their number who 
were engaged in splashing the water and 
throwing stones, frightened the fish and 
prevented their escape from ihe nets (Worsnop 
1897. 90-91^. 

George French Angas painted two men fishing 



with a seioe net at Second Valley, souih ol 
Adelaide, in 1844 (Angas 1847a: Plate XXli 
Cawthorne provides a similar scene, in 
watcrcolour, ai Second Valley, dated c.lS<*2 
(figure in Hemming & Jones 2000: 9), as does 
Sncll in his sketches, dated 14 July 1850, at Vorke 
Peninsula (SncI I 1849-59 1 1988- 128]). Eylmann 
(1908: 375 & Plate XXXIV fig. 3) published a 
drawing of two Ngarrindjen men using a long 
seine net. with a third man helping to drive the 
fish in. It is likely that several nets were 
sometimes connected together with supporting 
sticks. 14 In situations where nets were not 
available, branches could be used to drive the fish 
up on die beach (Angas 1847b. 112). A former 
colonist wrote to Tindale stating that in the Port 
Gerrnein area, north of Adelaide: 

On occasions a wall of prickly bushes was built 
in about ihree feet (one meltvj of water and 
rolled shorcwards, the ends converging to an arc. 
The fish which were impaled or caught were 
extracted by the pushers and thrown to others 
following behind (Parkes 1936, cited in Tindale 
1935-60: 48). 

The 'wall' of branches, comprised of saltbU'h 
and mangrove, was called wiiuia and required two 
people to roll it and two more for picking out the 
fish (Parkes 1936. cited in Tindale 1935-60- 96). 

Net fishing in fresh water required fewer people 
than sea fishing, with small nets used to catch fish 
living amongst reeds and logs. Nets weie essential 
items of Lower Murray material culture for fishing 
in the lagoons of the Lower Lakes, Coorone, 
Murray River and in the creeks of Ihe southern 
Fleurieu Peninsula. In the Encounter Bay area. 
Aboriginal women also caught tadpoles from 
elaypans with fine meshed nets and cooked them 
in large 'mutton-fish' shells (Worsnop 1897: 83). 

There was some degree of specialisation in the 
nets used by Lower Murray people. Harvey (1943: 
111) described three main lypes of nets used hy 
Yaraldi-speaking people of the Lower lakes. Nets 
with a small mesh, jairumi [pronounced 
'yatrumi'], were used when fishing for kanmuri 
(mullet, Aldrichetta forsteh). The nets with a 
larger mesh, neri, were used for lukkfri (bony 
bream. Nematulasa erebi). The big drum ne4s. 
dmgsri, were mainly for catching poiidi (Murray 
cod, MaccuHochella peeh) found under large logs 



'' In 1 '132 Tindale ( 1 953: J2) recorded from a 6unpwj| named Arthur Whirr dial Aboriginal people living along, rhc river had usco a double |minu?d 
wouden fishhook. Tindale (no dale., 'Milt-rum'. Siagt A. #3. Archives, South Australian Museum, Adelaide) also claimed dial Ihey used a flshTnj 
toggle made Irom bone as u form of fishhook. 

" This technique was also used in southern Yorkc Peninsula I Tindale 1036; 57) 



152 



PA CLARKE 



submerged in the river. 15 The drum nets were 
made with sticks in the form of a rectangular 
pouch (Harvey 1943: 111). 

The Tangani people of the Coorong used 
specialised nets, such as the jataruma 
[pronounced 'yataruma'] to catch mullet and the 
donggari to catch mulloway (Argyrosomus 
hololepidotus) and other big fish (Tindale 1934— 
37: 226). They also used the ngeiri net, which 
was made from the same sedges that were 
collected for basket and mat making. All nets used 
by Tangani people in the pre-European period 
were for hauling (Tindale 1934-37: 226). The 
Tangani used a hooped fishing net, termed 
kandarangku, with a mesh of 2.5 cm, for larger 
fish (Tindale 1951: 258). Apparently the term is a 
play on words meaning 'widow catcher', as 
kandari is applied to coarse rope and rangku 
refers to a widow. 16 Murray River people around 
Moorundie used the same type of net, 
kandenmku, to catch ponku (Murray cod). 17 The 
Kingston people used a net called pinang kanji to 
round up fish in shallow waters (Tindale 1931- 
34: 89). Angas provided a small watercolour 
painting of a pouch-like net 'for taking very small 
fish from Lake Bonney' (Angas 1847a: Plate LI) 
in the South East of South Australia. 

Aboriginal people in southern South Australia 
probably did not use set nets until well after 
European settlement. Tindale's main Lower 
Murray informant, Clarence Long (Milerum), 
claimed that set nets were rarely used in the 
Coorong when he was younger, and not at all 
before then (Tindale 1934-37: 226). This suggests 
that Europeans had introduced the practice to the 
local Aboriginal people. From available accounts, 
sinkers and floats were not used in association 
with net fishing during the pre-European period 
of southern South Australia. The record of Murray 
River people using clay from wasp nests to make 
sinkers for set nets is probably a post-European 
innovation.' 8 Tindale (1935-60: 17) suggested that 
vegetation placed along the top of the net to 
prevent fish from jumping out of the water might 
have helped keep the net vertical. 

It is likely that in the pre-European period most 



fishing nets in southern South Australia and 
Victoria were fabricated from two-ply string made 
from fibre obtained from the roots and stems of 
the bulrush (Typha species) (Beveridge 1883: 42; 
Krefft 1862-65: 361; Tindale 1935-60: 17; Zola 
& Gott 1992: 8-9, 62). Europeans also know this 
plant as flag, cumbungi and reed mace, whilst 
contemporary Ngarrindjeri people refer to it as 
manangkeri or manakeri (Clarke 2001b). It was a 
major food source, with the chewed remains of 
the roots being generally in good supply in 
Aboriginal camps (Clarke 1988: 69-70, 72; 
Tindale & Mountford 1936: 497). Tindale stated 
that: 

... mature rhizomes were chewed and fibres 
made into string, the chewing for this purpose 
being a duty of women. Older women's teeth 
were often worn to the gum line by their 
constant chewings." 

He also suggested that the proximity of bulrush 
grounds would have made a good riverside camp. 

In 1 894 an Aboriginal man from Point McLeay, 
Jacob Harris, provided a detailed written account 
of net making from 'rushes', which were probably 
bulrushes. He said that before Europeans arrived, 
Aboriginal people in the Lower Murray region 
used to: 

make our nets almost the same as the Europeans 
did, the meshes were the same, the only 
difference being that yours [Europeans] were 
made of twine etc., while ours from rushes. The 
rushes had to be steamed or cooked or whatever 
you like to call it, first a hole was dug, say a 
couple of ft. [ = 60 cm], then a fire was placed 
in it, after it had burned almost to ashes some 
damp or wet grass was put on the coals, then the 
rushes, so that the steam arising from the grass 
would steam or cook ... put in about evening and 
left until morning, it was then taken out and 
divided among the camp to chew or suck the 
juice out of it, which is very sweet, it was then 
made or twisted into a line for to be made into a 
net. ... the meshes were the same [as European 
nets]. 20 

Angas stated that the marine fishing nets were 
'composed of chewed fibres of reeds, rolled upon 



" Descendants from the Yaraldi and Maraura people from the Lower Lakes and Murray River regions made S.A. Museum drum net, A21338, in 

1934. 
'" Tindale (no date, 'Milerum'. Stage A, #3, Archives, South Australian Museum. Adelaide). 

17 Tindale. 'Ngaiawung vocabulary'. Archives, South Australian Museum, Adelaide. Also see Scott (cited Tindale 1951; 258). 
'* Tindale (no date. 'Milerum', Stage A, #3, Archives, South Australian Museum. Adelaide). 
rt Tindale (no date. 'Milerum', Stage A. #3, Archives, South Australian Museum, Adelaide). 
20 J. Harris letters (D65 10(1)14,15, Mortlock Library, Adelaide). Tindale (1934-37: 164) received a similar description of the process from his 

Ramindjeri informant, Reuben Walker. 



BARLY ABORIGINAL FISHING TECHNOLOGY 



153 



the thigh, and twisted into cord lor the purpose' 
(1847a: Plate XXI).' 1 Emu oil was sometimes 
rubbed into the fibre while it was being made into 
two-stranded cord (Tindale I93J-34: 60). Fibre 
treated in this manner makes the string more 
supple, while maintaining its strength and 
durability. Tindale (1951. 257) maintained that 
Aboriginal cordage from all areas of Australia was 
always composed of single or two-ply twists, 
never three or more. Nevertheless, there some 
were exceptions. For instance. West (1999: 20, 
23-28) describes the making of three-ply rope in 
Amhem Land and possibly Tasmania. 

After European settlement, changes in the 
Aboriginal diet would have led to a significant 
decrease in the availability of fibre from bulrush 
roots. Fibre used by Lower Murray people to make 
string for neis also came from an unidentified 
plant known in the local Aboriginal language as 
rulgoonowrie (MacPhcrson, no date. 10). A plant 
recorded as used for making Coorong nets was 
kuka (native flax, Linum rnarginale). 22 In 1932 an 
informant from the Maraura people of the Upper 
Murray of South Australia, Frank Fletcher, said 
that gill nets and drag nets were both made from 
the pungur rush (Tindale 1953: 23)." Harvey 
(1943: 109-110, 112) stated that the stems or 
jalken (knobby club rush, Isnlepis nodosa) were 
used for making nets." But field invesligaliom 
have indicated that the stems of this rush are pithy 
and not suitable For making siring or baskets. The 
stems of mangaiu (spiny sedge, Cyperui 
gymtwcaulos) would have been better: ihis 
species is still used by local Aboriginal people 
who make baskets and mats from it 25 Nets made 
from sedge stems were also made for dry uses, 
such as catching ducks.'"' 

The technique used lo make nets in southern 
South Australia has been variously described as 



'knotted netting' or 'knotted looping' (Davidscsn 
1933: 257-259, 269-272. figs 1, 9-10; West 
1999: 30-33, 49). The Tangani people called the 
mesh of the nei. nutndar. which reputedly meanl 
the- eyes'; the netting knot or lie was mulbakxm 
and the acl of making a ncl was lagulun (Tindale 
1934-37: 226). The last term is a reference to the 
threading motion of making the knot being like 
'spearing'." In the Ramindjeri language of 
Encounter Bay, i<ikk~in was recorded as spearing; 
weaving (as rushes for a basket)' (Meyer 1843: 
74). The mesh sizes for nets in southern Soul It 
Australia were variable, in the range 1-2.5 cm 
with the Kongarati Cave fragments, 2.5 cm 
recorded for the Coorong hoop net and 8 cm for 
netting wrapped around a desiccated body found 
at Fromm Landing, Murray River " 



Spear and Club Fishing 

During the day, men caught fish such as 
malawe (mulloway, Argyrosomus liolalepidotux) 
by standing motionless in the river or in their 
canoes (Hemming & Jones 2000: 9; Meyer 1846 
[1879. 192-193]: Smith 1930: 230-231; Sncll 
1849-59 [198S: 182]; Tindale 1934-37: 7| 
Fishers attracted the fish by their shadows and 
stabbed them with hand-held spears. Large speais 
made from Callitris pine witfi two to three 
wooden prongs lashed to the head were used for 
spearing mulloway and Murray cod J9 Being large, 
Uiese spears were also used as ptwling poles when 
fishers were travelling on hark canoes and rafts 
(Hemming & Jones 2000. 9). It is likely that the* 
spears were hidden near the area of Iheir use when 
the fishing season was over. In the Lower Lakes 
area fishermen frightened fish from their hiding 
places by a large noise, created by (hoisting one 



" A Hif'poning teteienie ('. Ariga.* ( IK<nh: Plate l.VJi. A-ii example ot a net made from Typho fibre is A2000. collected From the Lower Lakes district 

" Tiniliile luudiite. 'Milciuin. Stage A. iYLAkIuvcs. South Australian Museum. Adernldcl. 

" The tarn ptmjtur may relate to die iiunggunko specimen of Atnms specie* collected by Tindale from Swno Reach, 5 August l°M (AmWl.S ill 

Ausuolian Mu>*tin)> If so. »hen ihe plant retem=i* os for net-making was probably of the same specie*. Similarly, a related term /inn/icon, is a 

Western Victorian Aboriginal term tor die basket sedjte, Cam tfreticriuitj I Robsun 1 986). 
" Thil I'l.uil tyfcg bulntirly tfiuuil .is SqppU r»>'tostil. 

b I ■.' ample? of River Murray fishing nets made ftuut sedges (Cv/irnur species) in ilte South Australian Museum include specimens \ 1 7519. A2 13 IX 

ArtCiO. A,-4509t> Tberevm maiigant was recorded from Aboriginal sources in the IVKOs (CUitx 71X11 bf 
K Sec figure in Hemming * hints f20(Xl; 13). Smienhwaii (lyRh: .W-40) provided an overview of the use of nets lor catching waterfowl n 

Aboriginal Auin.'.l.n 
" In die Nganindjcii language of uV- Lower L*kcs, iWh/n was recorded to mean 'spearing' (Yatlop & Gnmwadc 1975: 33-M, 85-B6> 
■ Tindale U9S1: 2581 reported oo specimens A6509I. A25J5 1 and A20616. respectively Smyth ( I S7S 3OT-J90I discussed mesh sijzs hit ViMnnan 

neb. 
" In 1 1J6 Tindale collected a J I in-long fishing spear (Museum specimen AIGOOJ.t made of pine from the Coniong This specimen has two proof.* 

lushed in die head and was called /nuiln/uf/p&ri nuc-ivuvri in the Tangani language. In IR-I-* G French Angas painted a close-up of a lajge spear 

w i-t fltree prongs lashed on (Aitgas collection. Archive*. South Australian Museum, Adelaide — see Angas I S<* ?b Plate XKaij. This waimoloiu 

h ^produced in Hemming & Jones 121X10- y ). 



154 



PA CLARKE 



of these pronged spears into the water (Unaipon 
1924-25 [2001: 19]). The compressed air caught 
between the prongs rose to the surface with a loud 
report. There is an account of spear-fishing 
competitions held from November to April among 
the Piltindjeri group of Yaraldi-speaking people 
living along the southeastern shore of Lake 
Alexandrina, with the fish caught being presented 
to senior people in the community (Smith 1030: 
231-236. Unaipon 1924-25 [2001- I9-24I). In 
this 'nstance, it required skill in fish tracking, 
involving the detection of movements in 
pnndweed. reeds and water ripples. 

At night, a fire was used to attract fish to be 
siruck by spear or club (Angas 1847b: 112), 
Angas painted a club called a wadna, which was 
'used by the Port Lincoln natives, to throw at fish 
whilst swimming' (Angas 1847a, Plate LI). 
Similar beni elongated clubs from the Lower 
Murray are also In the Museum collection. Ciubs 
like these and described as boomerangs were 
reported as used along the Coorong (Olsen 1991: 
5). Sometimes a bark canoe was used as a 
platform to fish from, with a fire contained by a 
clay hearth in the middle that also served to cook 
ihe catcb (Angas 1847b: 54, 101, 107; Meyer 
1846 (1879. 193J) * Mobility across water was 
important for the fisherman in order to reach areas 
favoured by particular species of fish. Therefore, 
rafts made from reed stems (Phragmiies ausrralis) 
and grasstree flower-stalks (Xanthorrhoea 
species) would also have been important items of 
fishing material culture in the Lower Lakes and 
along the river lagoons." Although relatively 
common along the Murray River and in the Lower 
Lakes, bark canoes were rarely seen in the 
Coorong lagoon, and even then only for bringing 
in trade items. 33 Red gum trees (Eucalyptus 
lamaldulensis) required for bark are found only 
in permanent freshwater regions, such as along 
creeks in the southern Fleurieu Peninsula and 
along the Murray River. There is no evidence thai 
any form of watercraft was ever used for sea 
fishing, although short trips were made on reed 
raits or floats to nearby rocky islands in order to 
kill seals there (Tindale 1 94 1 • 241) 



Opportunistic H m v i -. 1 1 \< , 

There are documented cases of Aboriginal use 
of fish poisons in small lagoons of the upstream 
reaches of the Murray River bordering, northern 
Victoria (Curr 1883: 110). Nevertheless, Taplin 
doubled that Ngarrindjeri people possessed 
knowledge of the use of poisonous plants (Taplin 
1879: 47). When large numbers of fish died for 
natural reasons, such as changes in salinity in Ihe 
river or lake, Aboriginal people quickly gathered 
them (Eyre 1845, 2- 266) Before the construction 
of the barrages across the Murray Mouth 
separating the Coorong Irom Lake Alexandrina, 
elderly Aboriginal informants interviewed in the 
1980s claimed that there were sometimes rapid 
changes in the water from fresh to saline (Clarke 
2002). This tended to kill a large number offish, 
providing an abundani but temporary source of 
food. At other times, the incoming salt water 
drove certain species, such as Murray cod. 
upstream and suspended fishing in the lakes until 
it receded (Olsen 1991: 8; Taplin Journals: 
19 May 1860), Bony bream also die off seasonally 
in large numbers and, according to Ngarrindjeri 
informants in the 1980s, they had provided a 
temporary windfall of food (Clarke 2002). 

Fish-Traps and Enclosures 

Fish-traps and enclosures ranged from slight 
modifications to natural features of the landscape 
to special purpose-built structures (Mulvaney & 
Kammirtga 1999: 34-35). For example, in some 
areas Aboriginal people were able to use the 
narrowing of the channels between swamps as 
traps in which to place netted bags, tn his diary 
Wells records: 

Jusi now (July] there is fish lo be had out of ihe 
swamps, and we got one of the native females to 
make us a net. They are made of a long kind of 
soapy p|att [sic], bill flat and round, quite as 
targe as an umbrella top. tl is then doubled not 
unlike a huge ankle, boot, and at the two ends 
there is a small opening. It is then raised from 
ihe bottom in such a way thai the fish when 



" The Smith Australian Museum has a bark canoe iA6443) (mm Avoca SuUod on the Darling River dial contains an eaily 2<Nh century mock up 
made by museum artisans of Ihe firepUar lor display The original Fireplace was nude in a irml-linrd wooden container (A535S4) 

'' Jacob Hams, an Aboriginal man al Poini McLeary willing in 1891. claimed lhal Ihe use of ihe 'rude kind ol rati' made from reeds predated the use 
of bark canoes in Ihe Lower Murray (J Hams letters, Oti510(L.| 14,15. Muni.. I Library. AdcTMdc.l. The South Australian Museum holds rail 
soecimcns(AM6.12, A 14633) made by a Yaialili woman Amy Johnson, in 1930 

,: This statement derived from accounls by Mcimangk woman. Fanel Waison. and Tingaui man. Clarence l.ong (Milerum) (Tindale 1931-H 69. S7) 



EARLY ABORIGINAL FISHING TECHNOLOGY 



155 



caught cannot return. The net is then fastened 
with sticks in a narrow channel where one 
swamp runs into another and all fish going with 
the stream are caught. These fish the blacks call 
Coogolthcc. The nets arc laid overnight and by 
break of day next morning we have plenty 
brought us for breakfast." 

On the frontier of British colonisation, wild 
foods and Aboriginal labour were often crucial to 
Ihe survival of European settlers (Clarke 19%). 
Examples of more active manipulation of live 
environment were the construction of long 
trenches by Aboriginal people to concentrate fish, 
in the long term saving them much labour 
expenditure. For instance, Smith records that in 
the Murray region: 

In the low-lying country, alongside of the tivet. 
trenches are dug two or three hundred yards [180 
or 270 metres] long and from four to five feet 
|1.2 to 1.6 metres] deep. When the Murray 
becomes Hooded it overflows its banks to Ihe 
extent Of a mile or more [1 .6 kilometres or more] 
on each side, and frequently the Murray cod. the 
bream, the butterfish, and othet fish are living in 
this water When the waters become- low through 
evaporation and soakage the fish are easily 
caught ... They wade into the shallow water and 
stoop the fish into baskets made especially for 
this purpose (Smith I WO: 229). 

These channels were often relatively large 
modifications of the landscape that would have 
had a significant impact upon the local watcrtjltlf 
(Lourandos 1997 219-22). 

Another modification of the landscape for 
fishing was the building of stone and wooden 
fish-traps Some fish-traps, probably the 
predominantly wooden ones, were known by Ihe 
Ngarrindjeri name, ku:yitaypari y> The Yaraldi 
people if) the Lower Lakes did not make such 
structures, preferring to use fishing spears instead 
(Tindale 1931-34: 73-75) In the Coorong fish 
were harder to spear due to the prolific 
waterweed, so the Tangani relied heavily upon 
stone fish-traps, mlaipur (Tindale 1931-34; 73- 
75). Areas of shallow water in the Coorong, such 
as at the Yungurumbar crossing place, were 



considered good places for making such stone 
fish-traps (Tindale 1931-34: 71)." The Tangani 
generally placed their fish-traps made from 
limestone blocks along the landward shore of the 
Coorong (Tindale 1974: 61-62). This was 
presumably because the water here is shallow and 
covers a limestone shelf, in contrast to the 
seaward side of the Coorong lagoon, which is 
deep and sandy. Tindale provided a detailed 
record of the construction and use of fish-daps or. 
as his informants called them, ' fishing-stations' 
(Tindale 1931-34: 73-75, 1934-37: 5>.The fish- 
traps were generally made in the shape of a V. 
with one of the walls, termed itgatwngkure or 
ngalde. connecting with the bank. The wall that 
extended into the Coorong lagoon was sometimes 
up to 30 metres long The basket ware trap was 
placed at a gap in the base of the V. The species 
of fish seasonally caught in the traps included 
kcmgoldi (probably congollii, njindumi (an eel 
species), palengoi (unknown species), lapalap and 
therugarai (unknown fishes, not found in the 
Lower Lakes) and ten pateri ('.sand mullet') 
(Tindale. 1931-34: 75, 1934-37: 5). M No attempt 
was made to drive the fish in. relying solely on 
their natural movements. 

At Noonarocna, on the mainland side of the 
Coorong near Meningte. the lops of silted over 
fish-trap formations can still be seen, Aboriginal 
people interviewed by the present author in the 
early 1980s said that these fish-traps were 
naturally formed stone structures that were 
previously modified and maintained with stones 
and pieces of wood According to Ngarrindjkfl 
man Jack Koolmatric, mullet travel northwards up 
the Coorong during the day (Ely 1980). Upon 
reaching the trap, fish would swim into the wide 
mouth of a horseshoe-shaped line of rocks. The 
foundation of this was a natural reef of exposed 
rock with all gaps except one narrow exit shored 
up With logs and boulders. The mullet were forced 
to travel through this narrow exit where a net or 
basketry container put in their path would catch 
them. 

Elsewhere in southern Australia, some of the 
fish-iraps have been described in the literature as 



Well*. I t> July 1851 The specK5 of llsh relerred 10 hoc is probably kuniati icongoili. Ptnulnphrint urvilli) iCtvke 3001 b) 

Yattop ,& tirimwade (|y75: 55)iisi fotyin as 'rushes. siickC, and ku:yiroy/wi as 'fisb-vrap, bamei of slicks'. 

The Viingiinimbar crowing place is probably that between Rabbit. M toil and Sunggurungbar tprnnounced Y r unggiiningba(), a tut! on the 

Younghusband PtninsuU (see Tindalu 19?fi. fig U 

Ibr eel species referred !o here tS probably Atfful'f-t/ wstrutis. The pu/rngpl was said by T indole to be equivalent lo tlic YatulUi ptUliXS Tltik is 

piobabty fb&puUtnggi that Bemdt A Bcmdt (I9M3: 30M described ai 'mudfish' or •butittfish'- The topiitufi may possibly have been a sucies of 

Gulajuw, In Ute case of ien ptiter* or 'sand mullet'. Tindale (153 I-.I4: 74) said that its Yaraldi name is uria/iipr This i». probably ihe same word as 

the welappi' ot 'twtHei' recorded by Taplm 1,1879, 1311. EcVcrt & Robinson (L990: 19) consider thai Una nafbi to the 'jumping mullet' (i./;n 

Keyfenttj) 



156 



PA CLARKE 



weirs. 37 According to Edward J. Eyre, Aboriginal 
people seasonally gathered at the channels around 
Moorundie that connected the river fiats with the 
Murray River: 

making a weir across them with stakes and grass 
interwoven, [would] leave only one or two small 
openings for the stream to pass through. To these 
they attach bag nets, which receive all the fish 
that attempt to re-enter the river. The number 
procured in this way in a few hours is incredible. 
Large bodies of natives depend upon these weirs 
for their sole subsistence, for some time after the 
waters have commenced to recede (Eyre 1845, 
2: 253). 

This practice occurred during early December 
when the Murray River floods had already reached 
their highest point and were beginning to recede. 
Further south, in the shallow waters and swamps 
of Lake Frame near Burr Range, small fish were 
caught in weirs (Angas 1847b: 174). Also in the 
South East, near Rivoli Bay, Angas recorded: 

On some of the swamps the natives had built 
weirs of mud, like a dam wall, extending across 
from side to side, for the purpose of taking the 
very small mucilaginous fishes that abound in 
the water when these swamps are flooded 
(Angas 1847b: 155). 

In the pre-European period, Aboriginal 
modification of the riverine landscape for fishing 
must have been considerable. It was oral history 
among old river boat captains in the 1980s that 
before the lock system was introduced, remains of 
Aboriginal built fish-traps were known hazards to 
paddle-steamers when the water level was low (T. 
Sim, pers. comm.). Hahn recorded that in the 
Hahndorf area of the Mount Lofty Ranges, the 
Aboriginal people would: 

build a dam into the river, high enough to let 
about a foot [31 centimetres] of water stream 
over it. Because of this dam, the fish in their run 
must come close to the surface of the water, 
where the savages stand in readiness to spear 
them (Hahn 1838-39 [1964: 133]). 



In coastal zones weirs of brushwood 
constructed at mouths of creeks caught fish left by 
receding tides (Angas 1847b: 112). Some of the 
trenches, traps and weirs were designed to catch 
bait for much larger fish. For example, near 
Martin Well on the Coorong, drains 100 m in 
length were constructed by Aboriginal people to 
catch small fish, called lap-lap, for bait (Worsnop 
1897: 106). 3S These were netted in fine close mesh 
nets. These weir and trench constructions appear 
similar in design to the much larger earthworks at 
Lake Condah in western Victoria. 39 

Outside the Lower Murray, but elsewhere in 
southern South Australia, fish-traps have been 
recorded in the Port Lincoln estuarine area 
(Martin 1988; Mountford 1939), as well as at the 
mouth of Pedlar Creek south of Adelaide 
(Stapleton & Mountford, no date). In the case of 
the latter, the Tangani man, Milerum, knew of 
these traps as he and his family camped there, 
when he was a child, on their way to Adelaide via 
the coast. 40 There were also fish-traps made from 
large water-worn boulders at Moana Cove, also 
south of Adelaide, but these were in poor 
condition when Tindale inspected them in the 
1920s. 4 ' Some traps could be dragged out of the 
water when fish were not required. For instance, 
Charles Sturt stated that on his 1830 expedition 
down the Murray River he: 

observed some cradles, or wicker frames, placed 
below high water-mark, that were each guarded 
by two natives, who threatened us violently as 
we approached. In running along the land, the 
stench from them plainly indicated what they 
were which these poor creatures were so 
anxiously watching (Sturt 1833, 2: 165). 

Small dams or 'pounds' might not only be 
structures for catching fish, but were also used to 
keep part of the catch alive for future use. At the 
Point McLeay Aboriginal Mission, 'fish pounds' 
made of stakes were in common use by Aboriginal 
people during George Taplin's period there 
(1859-79). 42 According to Lower Murray man 



" For example, Beveridge (1883: 48); Curr (1883: 110) and Pretty et al (1983: 119). Tindale listed 'fish weir (set in creek)' as ake in a Murray River 

dialect (Tindale, 'Ngaiawung vocabulary'. Archives, South Australian Museum, Adelaide). 
u This fish, lap-lap, was possibly a species of Galaxias, Lapps Lapps was recorded to mean 'small fish' in the Booandik language (Smith 1880: 3). 

Wells (1852-55) listed lap-lap as 'very small fish', Tindale (1931-34: 74) said that lapalaf) were a Coorong fish species not known in the Lower 

Lakes. 
" For a description of the Lake Condah eel-traps, see Courts et al (1978); Flood (1995: 240-245; 1999: 216-220); Hemming (1985); Robson (1986) 

and Worsnop (1897: 104-106). The S.A. Museum has a basketry trap, A6431, collected from Lake Condah. c.1910 (see figure in Hemming & 

Jones 2000: 9). 
" Tindale (no date. 'Milerum'. Stage A. #3, Archives. South Australian Museum, Adelaide). 
41 Tindale (no date, 'Milerum'. Stage A, #1, Archives, South Australian Museum, Adelaide). 
" The 'fish pounds' were recorded by Taplin on the shore of Lake Alexandria (Journals, 11-12 October 1859; 8 November 1859; II November 

1859; 28 November 1859; 10 January I860; 21 November 1861; 20 March 1862). 



EARLY ABORIGINAL FISHING TECHNOLOGY 



157 



Lindsay Wilson, interviewed by the present authoi 
in the early 1990s, the practice of fish storage in 
wooden enclosures in the lake at Point McLeay 
continued until ihc 1930s, Dunne the early 
twentieth century along the Coorong, European 
fishermen kept alive the bream ihcy had caught by 
putting them in 'pounds' made of tea-tree stakes, 
until the weather was suitable fur transporting 
them to the market (Lvans 1 99 1 ; 40) 



Cu.U'Ral Aspects 

Aboriginal people along the Murray River 
considered that their Dreaming Ancestors created 
the wetlands (hat provided them with abundant 
sources of food. For instance, Natone, an 
Aboriginal man from the South Australian section 
of the Murray River, claimed dial a blind woman, 
Noreela, had created the environs of the river 
(Bellcnainbers 1931. 112, 125). Starting from 
Lake Victoria, Norecla. with two young children 
tu guide her, made the river by driving back the 
Sea. She travelled like a 'drunken bee' her 
meandering course meant that lhe river was very 
long This lengthening of her journey was 
considered greatly desirable, as it increased the 
number of fishing and hunting grounds, with a 
lagcvn at each elbow. The fossils jutting out of 
cliffs along the Murray River were said by 
Aboriginal people to be the remains of fish killed 
and eaten by Norecla and her children. Another 
account, still told by Ngarrindjcn people in the 
1980s, was of a Thukabi Ancestor, who was .i 
large river turtle that came down lhe Murray 
River: 

Thukabi |u large river lunle) came down (from 
the Darling district] through rhe desert looking 
for a place to lay its eggs. As she went, lira drag 
of its tail made the river its flipper carved oul 
lhe lagoons and banks. You cui sec where it 
wcnl. When il got 10 the laHe, it pushed itself 
into the sea (H Rankine cited Clarke 1994. 
114). 

In all these accounts, lhe present course of the 
Murray River is explained by the actions of 
Ancestral Beings. 4 ' 



In the Ngarrindjcn Dreaming of the Lower 
Murray, Ngurundcri chased a large Murray cod. 
Pondi, down the Murray River, widening its banks 
in the process, until it was eventually speared near 
the Murray Mouth (Berndt & Berndt 1993: 224, 
Clarke 1995: 148-149; Hemming &. Jones 
20Q0). 44 Whilst Ngurunderi was at the Murray 
River entrance into Lake Alexandrina, a waicr 
spirit known as the Mulgycwonk tore holes in his 
nets, which prevented him fishing for his family 
(Tindale & Pretty 1980: 50). There are accounts 
of Aboriginal legends from further upstream in 
Murray-Darling Basin where the dreaded river 
spirit who fed on boys was a large Murray cod 
(Sinclair 2001: 120-121, 252). Ngurunden made 
the fishing grounds and lookouts for the Lower 
Murray people, which is told in song (Tindale 
1931-34- 259). 

Fish species also appear elsewhere in Lower 
Murray mythology. For instance, Yamakawi 
(Shark Ancestor) had a prominent role in the 
Kondoli (Whale Ancestor) Dreaming of 
Encounter Bay, which explained the origin of fire 
(Clarke 2001a: 24-25). There is also a Dreaming 
myth relating predominantly to freshwater fishing 
technology, which took place in the region from 
Lake Hindmarsh in western Victoria to Lake 
Alesandrina in the Lower Murray (Harvey 
1939ms, 1943, Tindale 1934-37: 65-69). In this 
account the drum-nets of the pelicans were 
transformed into large pouches under their bills. 
In the Dreaming of Limi < = Lime), die love of 
eating fish is involved in explaining the creation 
of the Inman and Hindmarsh Rivers of the 
southern Fleurieu Peninsula (Bcmdt & Berndt 
1993: 311; Meyer 1846 [LS73: 202|). J5 The 
Skyworld landscape contains a fish too, with a 
constellation of stars - being Nunganari, lhe 
Stingray (Berndl & Berndt 1993: 164, fig. 25; 
CJarke 1997: 131). 4 * 

Economically important fish species were 
considered by the Lower Murray people to have 
been created by Ancestors during the creation 
period. For instance, in a Ramindjeri account of 
the Kondole (Whale) Dreaming, lhe Kuratje and 
Kanmari Ancestors became small fish when ihcy 
ran in the sea: 



"' 1 rom the larc 1980s, an animated version of Ihis mylh was given nl Ihc Sipiiil ROioJ Mu<eitm, Goolwa. 

" Sinclair (21X11: 120) speculated dial the relatively large art lhal some cod grow to. up 10 1.8 metres, enhanced Abm1gin.1t beltcls or die beasi '« 

mythical ,|Ualiut'5- 
" Berndt & Berndi ( I W 111) suggested thai Limi was a Stingray or Carpel SbSifc Ancestor. Meyer 1 1 MS pan ? "5 1 lilted Lime/ a* a 'kind of 

ie. t r f.iini .vid hit family were eventually transformed inlo a group ul pttMJrrSlU lock! in lhe ;ec, al Victor Harbor near ihc Ltlull (Meyer 1846 

[1879- 202J). 
"' According to Tindale Inn dale. Milemm , Stage A, #1. Archives. Suutli Australian Museum. Adelaide), the Tangam considered lhal there were 

seven stars shaped like a stingray. 



158 



PA CLARKE 



The latter was dressed in a good kangaroo skin, 
and the former only a mat made of seaweed, 
which is the reason, diey say, that the kanmari 
[mullet, AUrichetta ft/rsicri] contains a great 
deal of oil under the skin, while the kuratje 
[Western Australian salmon. Arripb truiiaceux] 
is dry and wilhout (at (Meyer 1846 [1879: 
2QS3)> 

In other Dreaming accounts, the Murray cod, 
1'ondi, was cut into smaller pieces by either 
Ngumnderi or Waiyungari, depending on the myth 
version, and thrown back into the water to become 
different ivpes of fish (Clarke 1995: 148; »999b. 
53-S4). 

The Raniindjeri version of the Ngunindcri myth 
epic, recorded from Reuben Walker in 1935, starts 
from a large Sake near mountains somewhere to 
the northeast of the Lower Murray and involves a 
large fish which is not a Murray cod- 

Ngurunderi came to the Lake and speared the 
fish which, made off with it at a great speed and 
cut a deep track right down to Lake Alexandrma. 
Ngurunderi followed and noticed that the river 
thus formed was without fish. So he stood on the 
bank and broke die bark of the red gum tree 
(H'Urf) lip into shreds & threw it into the water 
and said; you arc Murray end. This must be true 
because, when you cut open a Murray cod you 
can see a tree like mark of blood vessels on the 
walls of its body, it is just like a gumtree 
Ngurunderi then threw mpujulanki bush (Native 
wild grape'.' Cryjiiandra hisptdula) and this 
became cattish (Tamianus tandanus). He threw 
in peppermint gum bark (lentumi) and it became 
the ijiri (Jerapon bidyanus Mitchell). After he- 
had made all ihe best fishes he threw in a piece 
of refuse and it became the Bony bream (tulcari, 
Fluviuloau richardsani (Caselman)).*" 

In relation to the blood vessels, it is worth 
noting that along the Murray European fishers 
have an analogous belief which interprets the tree- 
like markings on cod swim bladders as 'skin 
maps' that show the birth place of the individual 
Osh (Sinclair 2001: 124-127), Tindalc recorded 
from a European settler the following Dreaming 
account involving Matumen who: 

... chased a large fish from the sea entrance of 
the Murray Mouth up through the Lake until he 
caught it out off Poltallock IPoltaloch Station) 
where he killed it and pulled it to pieces and 
ihrcw the pieces all about and they turned into 
small fishes and that is how the different sorts of 



fish came (RD Anderson 1934 [cited in Tindalc 
1934-37: J75)). 

The Ancestor mentioned here is probably 
Matamai, who was Ngurunderi's son (Clarke 
1995: 146) The theme ol an Ancestor tearing up 
large beings to make many smaller species also 
occurs in relation to kangaroos according to one 
account of the Waiyungan mythology (Clarke 
1999b: 54). 

Aboriginal people considered themselves as 
having a role in the continuation and wellbemg of 
i heir environmental resources. Ceremonies were 
sometimes performed in order to increase fish 
supply. For instance. Howitt relates: 

There is a spot at Lake Victoria | = Lake 
Alcxandrinal, in the Narrinyeri [ = Ngamndjenj 
country, where when the Water is, at long 
intervals, exceptionally low, it causes a tree- 
shimp to beeome visible. This is in charge of i 
family, and it is the duty of one of the men !o 
anoint it with grease and red ochre The reason 
for this is lhal they believe that if ir is nor done 
the lake would dry up arid the supply of tish be 
lessened. This duty is hereditary from father to 
son (Howitt 1 904; 3W-400). 

The illicit involvement between Waiyungari and 
Nepeli's. wives in the Ngarrindjeri Dreaming was 
perceived as the cause of poor fishing in early 
spring each year. The arrival of (he Young Men 
(Orion) and the Women (Pleiades) constellations 
in September was considered to help turn tins 
around (Berndt & Berndt 1993: 164: Claikc 
1999b: 57). The flowering of certain plants may 
also have been an indication of the arrival in the 
Lower Murray ol certain species of fish. This was 
the case at Marion Bay in Yorke Peninsula, where 
Ihe prolific (lowering of tea-trees was a sign to 
the Narangga people that the mullet fish were 
soon (o come in large numbers." Here, il was 
claimed (hat initiation ceremonies were held then 
to take advantage of this seasonally abundant food 
source. 

The abundance of fish would have allowed for 
a larger Aboriginal population in the Murray 
Basin in comparison to the surrounding regions. 
The Lower Murray was particularly rich in fish 
resources, involving marine, estuarinc and 
freshwater species (Eckcrt &. Robinson 1990; 
Evans 1991, Glover 1983; Olsen 1991; Sim et al 
2000). Angas said 'The Milrnenduras subsist 



Idcniilicaiion ol ihese lish n.crncs is given By &rktn $ Rohinsun I l l J90: 19-20,1. Nole lhal kiirmve h cquivalcni to tamu.- 

Tindalc < I9JIM.Z' ! 1 1). [Italic* by. tilt present oulhor,J 

Reminiscences recorded hy F. DtVMt in file Mail newspaper. Adelaide on 25 March 1952. 



EARLY ABORIGINAL FISHING TECHNOLOGY 



19 



chiefly on fish, and though extremely wild and 
treacherous, present some of the best specimens 
of the Aboriginal Inhabitants, as regards physical 
appearance' (Angas 1847a: Plate XI). Angas also 
commented that ' On the S. E. coast and along the 
shores of the Murray and Lakes Alexandnna and 
Albert the natives live chiefly upon fish, and 
waterfowl' (Angas 1847a: General Remarks) 3 " 
Aboriginal people considered that some of the 
spirits with whom they shared the landscape also 
liked eating fish. For example. Lower Murray 
people believed that the dreaded river spint, the 
Mulgycwonk, was attracted to the smeJl of fish 
and once captured a young boy who was washing 
lish oil from his hands on the edge of the lake 
(Clarke 1999a; 157; Harvey 1939ms) 

Profit the daily account available in Taphn's 
Journals, it is clear that fish, termed tiwme in 
general, were a favourite food item »n the Lower 
Murray region. NgaTrindjcri people considered 
that the dominance of fish in their diet set them 
apart from at least some of their neighbours They 
gave cultural significance to the fact thai when 
ilieir babies tried to speak, their first word was 
mam tTaplm Journals: 10 October 1861). Adults 
proudly interpreted this as the infant's desire to 
eat fi«.h. The fish entrails, ngarakuni. were 
considered good eating, being grilled over the 
coals with the edges of the fillet curled up to catch 
the juices. - Special slicks, wtinupi, were used as 
fire tongs to remove food, such as fish, from hot 
coals." Fish remains are often found in Aboriginal 
middens on ihe banks of the. Lower Lakes and 
Murray Rher fLuebhers I97S, 1981, 19S2; Pretty 
etal 1983: 1 1 7- U 8; Tindale 1930-52:67) 

Some Aboriginal groups in southern South 
Australia were noted by their neighbours for 
having a diet dominated by flab. This is shown bj 
a recorded remark by Parnkalla people of 
northeastern Eyre Peninsula that their Port Lincoln 
neighbours, the Nauo, had 'an offensive breath, 
being fish eaters' (Schcirmanri 1844, 1. 7). In the 
Lower Murray, Murray cod oil was rubbed on 
initiates (Tindale 1930-52: 139). The Tangant 
people of the Coorong had a song 'ridiculing men 



who refused to lend their fishing net" (Tindai:- 
1934-37: 267). 

The material culture of the Lower Murray 
people reflected their fishing background. Old 
fishing nets were used in the Murray Basin for 
wrapping human bodies that rnd already been 
desiccated CSheard et al 1927- 173. Tindale I95J: 
258; Tindale & Mountford 1936: 495, 499), In the 
Lower Murray, fishing nets were often among the 
personal items placed in burial bundles lodged on 
tree platforms (Hackett 1915: 29). Stingray tail 
barbs or 'nails' were used in ngildjeri sorcery 
(Bemdt & Berndt 1993: 260) In the Murray River 
area, Tindale recorded the medicinal use by 
Nganguruku people of Eucalyptus leaves, tindunj. 
with fish fat. Apparently ihey would 'inluse 
leaves in bark dish over hot ashes, mix with fish 
fat (liver) for colds'.-- Angas painted an object 
that he described as 'Kuikoonga - Bones of a fish 
found in the Murray, wom as a head ornament, in 
the same manner as the teeth of the kangaroo' 
(Angas 1847a: Plate XXX). String-games or 'cats- 
cradles' played by Lower Murray and South East 
women and children often featured 'fishing nets' 
(Tindale 1931-34: S8). M 

The wide distribution of recorded modifications 
to the landscape Id assist in eapiuring or storing 
fish in the Lower Murray and neighbouring 
coastal and riverine areas indicates that their use, 
in prc-European times, was a major subsistence 
strategy. Lourandos has described some- 
earthworks by Aboriginal people in temperate* 
southern Australia as artificial drainage systems 
operating to flush fish from swamps into channels 
set with traps (Lourandos 1997. 219-221. 22 rj 
This form of swamp management coped with 
excess water during floods and helped retain water 
in times of drought. This was part of a subsistence 
pattern that allowed for a larger and semi- 
sedentary human population, in comparison in 
other regions. The material culture and diet of 
Aboriginal people living in the south would 
therefore have significantly differed from groups 
to the north and particularly those situated some 
distance away from major bodies of water. Fish 



iupptnifg ;eferencei «e Angus (1847b: F'lalcs IX & XXV). 
" HaivcyM)939[ns> and Tindale (, 1930-52; MH-i-W) iktu: hed the manner in which fish weft cm up »ndh»>t|sivcu the Ydialdi names foi die piem. 
'■' l i/idalc &. Mounl/ord (19V., 49f,| Sit photopaph In Mrmming & Jones (2000: 17i. 
' M ti 1 ind-.ilc specimens (tVff|*7S - tiUriAftfRUJ <**IiJ! AfiSjtiS - £ fr/rtw/o, A7SES - unknown £m u/vj.ia.. spu-tan.) collected 9 mites nonh nt 

wur Reach 5 Aueij'l 1964 Clirite (l"**9 : 3) ha; a similar record relating to mullowav live) 
'■* In 1930 Tindale collected a stnng-pame iAU'icCt. called 'fishing net", from Amy lohnson (YajaJdi people. Lower Lakes! A stnite-gimie 

I Vj6733), described asaTaxtgam 'lish net game' from the Coorong was possibly a replica made hy Tindjl? in |93d (sec Hemmine & Jones 200<>: 

|9) In WO O.S Davidson and N B Tindale collected a( Swan Rca,* a stnng-ramt (Al4'r58), called a 'net', mode by Jerry Mason (Yiiamka 

people. Murray Riverl- 



160 



PA CLARKE 



storage in pounds and mud pools was a pre- 
European Aboriginal practice for managing the 
windfall/drought situation of food gathering. 
Another Lower Murray technique to extend the 
use of fish food in pre-European times was to dry 
fish on racks (Berndt & Berndt 1951: 29). 
Because of the highly seasonal nature of fishing, 
it is likely that some of the stone and wood 
structures found in southeastern Australia, 
identified by archaeologists as fish or eel-traps, 
would be better described as fish pounds, being 
for storage rather than capture. 

Among the Lower Murray people, many of their 
totemic familiars, the ngailji, were species of fish 
(Berndt & Berndt 1993: 306-312). This was also 
the case for totemic groups along the Murray 
River (Tindale 1953: 37, 49). In their daily life, 
menstruating women in the Lower Lakes and 
Murray River areas were forbidden to eat fish 
(Berndt & Berndt 1993: 124-126, 141; Eyre 1845, 
2: 295; Harvey 1939ms). These women were not 
allowed to go near the water at all, as it was 
considered that the success of men fishing would 
be spoiled. On one occasion in the past, a large 
incursion of seawater brought a considerable 
number of poronti (sea mullet, Mugil cephalus) 
into the Coorong lagoon (Tindale 1931-34: 1 19). 
In spite of their abundance, amongst the Tangani 
people only old men were allowed to eat them. 
The mulloway would swim into the Coorong 
lagoon to spawn under the limestone cliffs 
(Tindale 1931-34: 119). In this condition only old 
men were allowed to spear them. Similarly, if 
young men ate pelenggeri fish (unknown species), 
they would prematurely become baldheaded and 
grey (Tindale 1934-37:39). 

In the Lower Lakes no Yaraldi women, except 
the elderly, were allowed to eat catfish (Harvey 
1939ms). Similarly, young Yaraldi girls were not 
allowed to eat big-bodied kai.kuanggi 
('freshwater bream' - possibly black bream, 
Acanthopagrus butcheri). Women during 
menstruation were not allowed to go near water or 
to eat fish caught with nets, as the Yaraldi 
believed it would drive the fish away. In recorded 
versions of the Ngurunderi Dreaming, his fleeing 
wives cooked and ate tukkeri (bony bream) fish. 55 
The importance of this particular episode is 
explained in various ways, depending on the 
particular account. These are that the wives were 



breaking a food prohibition by eating this fish 
species; that the fat exploded in the fish, making 
them sacred to Ngurunderi; and that the wives 
incurred Ngurunderi's wrath by giving him the 
smaller of the two fish they had cooked. The 
unifying theme in all versions is that this fish was 
not to be eaten by women. 

In the Murray River region more restrictions 
were placed upon females until past the age of 
child bearing than upon males of the same age 
(Eyre 1845, 2: 293). Teichelmann stated that in 
southern South Australia, food prohibitions were 
such that women with children were prohibited 
from eating certain food and that they generally 
lived upon vegetables (Teichelmann 1841: 7). The 
categories of people with least amount of food 
prohibitions appear to have been infants and the 
elderly. Eyre said that in the Murray River area 
'No restrictions are placed upon very small 
children of either sex, a portion being given to 
them of whatever food their parents may have. 
About nine or ten years appears to be the age at 
which limitations commence' (Eyre 1 845, 2: 293- 
295). He also stated that old men and women were 
able to eat most things. All prohibitions would 
have been based on cultural logic, even if the 
reasons were obscured from those practising the 
ritual. People generally excluded from 
prohibitions were those not sexually or 
economically active. It was the power to produce, 
either physically or spiritually, at particular life 
stages that was perceived as making people 
sensitive to influences potentially harmful to 
group harmony. In a sense, an individual's 
position in the society could be defined by what 
the person could eat and what economic activities 
they could engage in. The major categories 
determining prohibitions appear to have been age, 
gender and initiation status. 

Aboriginal people could, at certain times, 
exercise a degree of choice in the food they lived 
on. For instance, some Lower Murray animal 
foods such as fish, emu and kangaroo meat were 
highly favoured foods when available. Yet 
vegetable foods such as roots were probably the 
mainstay when meat was not easily obtainable 
(Clarke 1988: 73-74). A report from the 
Adelaide-based Statistical Society in 1842 
illustrates the seasonality of Aboriginal food in 
the southern areas. 56 The report notes that in 



Accounts summarised from Bemdt & Bemdt (1993: 224, 435), Clarke (1995: 149, 1999b: 54) and Tindale (1934-37: 285). 

'Transactions of the Statistical Society. Report on the Aborigines of South Australia.' Register newspaper, 8 January 1842. Thomson (1939) also 

considered the seasonal aspects of Aboriginal culture. 



EARLY ABORIGINAL FISHING TECHNOLOGY 



161 



spring mainly vegetables and grubs were eaten. 
With the commencement of summer, fish were 
obtained, as were kangaroos, emus, U/ards and 
the eggs and yuung of birds. During the hottest 
pait of the year possums and acacia gum were 
procured, while in autumn berries and nectar were 
.available. In the winter a variety ol roots were 
consumed, as were possums and other animals. 

The coaslal zones ol southern South Australia 
were rich in natural resources, particularly food 
such as fish, molluscs and coastal berries. Meat 
from occasional whale strandings was also an 
attractive coastal food source (Clarke 2001a) 
Although many of these foods were available for 
the greatei part of the year, the onset of winter 
made the coast a harsh zone in which 10 live. 
Partly for this reason, 'salt water" Aboriginal 
groups in southern South Australia would have 
moved according lo season betweeo inland and 
the coast. Stable isoiope analysis of human hone 
indicate* thai 'sail waler' groups did not penetrate 
up the Murray River beyond lhe boundaries of the 
Lower Murray cultural region (Pate 1997, 1998, 
2000). The pattern in the Adelaide region was a 
general movement away from the coast in late 
autumn, so that more substantia] shelters could be 
built in the protected Mount Lofty Ranges 
foothills (Clarke 199 1 - 58-59; Ellis 1976: I lo- 
ll 7; Ross 1984- 5; Tindale 1974: 60-61). The 
historical record shows similar earJy patients for 
coastal groups from the Lower South East of 
South Australia (Foster 1983: 23-43). Seasonal 
movements, although an aspect of the Aboriginal 
relationship to the physical environment, are 
essentially dictated by the 'cultural landscape 1 . 

A seasonal population movement occurred 
among at least some Lower Murray Aboriginal 
groups, (n winter the 'salt water' Tangani people 
camped along the mainland side of the Coorong 
lagoon, where Firewood was plentiful and shelter 
from weather available. 5 ' The fish-traps 
maintained there, where the water is shallow, 
provided a reliable source ot food. During 
summer these Tangani people camped on the 
Yniirighusband Peninsula between the Coorong 
and the Southern Ocean, giving them easy access 
lo coastal Poods such as marine Fish and berries. 
I'll, re were also political teasons for ihc 
movements, with the actions of neighbours 
impacting on Lower Murray groups. For example. 
one of lhe disadvantages of camping on lhe 
mainland side of lhe Coorong was thai Here the 



Tangani were open to attack from lhe Ngarkal 
people, who normally ranged in mallee areas i< ■ 
the east of the Lower Murray. During harsh 
summers the Ngarkat people were forced towards 
the Murray River and Lower Lakes when their 
water supplies dwindled, but the Tangani 
considered that they were not likely to be attacked 
during the winter, The locations of most Lower 
Murray seasonal camps are not known, although 
more archaeological research may improve our 
present knowledge. For 'fresh water' Lowci 
Murray people, such as the Yaraldi-speaking 
groups living along the edge of Lake Albert and 
Lake Alexandria, lhe yearly movement was 
probably from the Iakcshore lo nearby inland 
areas, in order to maximise food supplies and 
comfort (see Fig. 2). During warmer months 
lakeside camps were cooler as well as being close 
to freshwater food sources In contrast, during 
winter, campsites in close proximity to large 
bodies of water were more exposed to cold 
weather. Back from the lakeside, the forests 
piovided natural windbreaks and had more 
Firewood and hut building materials available. The 
prominence of aquatic technology used by the 
Lower Murray people would have given them 
little interest in the remote and harsh inline! 
regions. 



Conclusion 

Resource usage by early Lower Murray 
Aboriginal people was a function of lhe broader 
environmental and regional patterning of lhe 
landscape. Aboriginal people in the Lower Murray 
were not randomly dispersed over the landscape; 
Ngarrjndjeri people were restricted to Die riparian/ 
marine areas that were consistent with their 
material culture. They were, among other things, 
regionally organised according to their perception 
arid use of the natural resources. The Lower 
Murray people considered their relationship with 
the environment to be an active one. Not only did 
Ihey physically manipulate their resources, they 
also considered ihcmsclves ro be influenced and 
organised by the environment The material 
culture of the Lower Murray people, although 
having some aspects in common wilh water-based 
subsistence cultures in the South East and the 
Murray River regions, was distincrive. They were 
largely a cultural group confined to the npaiiaiv' 



TinJdc 119.18- II. 1974. f, I -Ml. 



162 



PA CLARKE 




I I - OPEN SCRUB 



COASTAL 
SUCCESSION 
Kingston! 



Coastal Groups 



Inland Groups 




Winter camps 



COASTAL 
SUCCESSION 



WOODLAND 



SOUTHERN 
OCEAN 



COORONG 




LAKE 
ALBERT 



FIGURE 2. Pre-European vegetation of southern South Australia (after Boomsma & Lewis 1980: map) and 
Aboriginal camping zones in the Lower Murray. 



EARLY ABORIGINAL FISHING TECHNOLOGY 



Lti 



coastal habitats of the southern coastal region of 
South Australia. Their hunting and gathering 
practices help to define this cultural region. 
Approaches in cultural geography, focusing on the 
cultural construction and perception of the 
landscape, are well suited to the study of 
Aboriginal environmental knowledge, 



At KNOWIJIDtiMFVrS 

A draft of this paper appeared in the author's PhD 
thesis, which was supervised hy C Anderson, P StfQites 
and K Garbett T Sim checked Ihe scientific names of 
the fish species. 



Refbri 



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Melbourne. 



FOSSIL LIZARDS FROM THE PLIOCENE CHINCHILLA LOCAL FAUNA, 
QUEENSLAND, WITH DESCRIPTION OF A NEW SPECIES 

M. N. Hutchinson & B. S. Mackness 



Summary 

The lizard fauna of the Middle Pliocene Chinchilla Local Fauna consists of members of the 
Gekkonidae, Agamidae, Varanidae and Scincidae. A new species of the scincid lizard genus Tiliqua 
is described, based on a complete right dentary. The new species combines relatively unspecialised 
dentition, most similar to that of T. nigrolutea, with large size and relatively gracile proportions, 
with a relatively slender coronoid process and deeply concave margin between the coronoid and 
angular processes. An additional scincid dentary is probably referable to the genus 
Cyclodomorphus. Cranial remains of a species of the gekkonid genus Diplodactylis are described, 
representing a group of species within Diplodactylus that have a vestigial jugal. An agamid dentary 
and two varanids of differing sizes are also described. 



FOSSIL LIZARDS FROM THE PLIOCENE CHINCHILLA LOCAL FAUNA, QUEENSLAND, 
WITH DESCRIPTION OF A NEW SPECIES 



MN HUTCHINSON & BS MACKNESS 



HUTCHINSON, MN & MACKNESS, BS 2002. Fossil lizards Imm Ihe Pliocene Chinchilla 
Local Fauna, Queensland, wilh a description of a new species. Records of the South Australian 
Museum 35(2): 169-184. 

The luard fauna of ihe Middle Pliocene Chinchilla Local Fauna consists of members of the 
Gekkonidae, Agamtdae. Varanidac and Sctncidae. A new species of the scincid lizard genus 
TiliqiM is described, based on a complete right dentary. The new species combines relatively 
unspecialiscd dentition, most similar to that of T. ntgmluiea. with large size and relatively 
gracile proportions, wilh a relatively slender coronoid process and deeply concave margin 
between the coronoid and angular processes. An additional scincid dentary is probably referable 
to the genus Cyclodomorphus. Cranial remains of a species of the gekkonid genus 
Diplodactylus are described, representing a group of species within Diplodactylus that have a 
vestigial jugal. An agamid dentary and two varanids of differing sizes arc also described. 

Mark N Hutchinson, Department of Heipetology, South Australian Museum, North Terrace. 
Adelaide, South Australia 5000. Brian S Mackness*. School of Biological Sciences, University 
of New Souih Wales, Kensington, New South Wales 2052. *Current address: PO Box 560, 
Beerwah, Queensland 45 19, Australia Manuscript received 17 April 2002. 



The Chinchilla Sand was named by Woods 
(I960) tor a sequence of weakly consolidated grey 
to yellowish and light brown sands, fertugitrised 
heterogeneous conglomerates, grits, sandy clay 
and clays. These outcrops range from shallow 
beds to sections several metres deep along a 
40 km stretch of the Condaminc River valley, 
approximately 240 km ENE of Brisbane. 

The Chinchilla Sand has yielded a diverse array 
of fossil remains, the Chinchilla Local Fauna 
(Mackness, Wilkinson & Wilkinson 1999). 
Mammals have been die best studied croup and 
the fauna recovered comprises macropodids 
(Bartholomai 1963, 1966, 1967. 1973, 1975. 
1976; Flannery & Archer 1983), phascolarcods 
(Archer 1977). tltylacoleomds (Archer & Dawsnn 
1982'. Bartholomai 1962, Woods 1956). 
pcramelids (Mackness ct al 2000), dasyurids 
(Archer 1982; Bartholoimii 1971; Dawson 1982; 
Wroe <fe Meekness 1998. 2000a, 2000b), a 
thylacinc (Mackness. et al submitted), murid 
rodents (Godthelp 1990) and a molossid bat 
(Hand et al 1999). At least three lineages of birds 
ate also known, an emu, a rail and a duck (Olson 
1975. 1977; Patterson & Rich 1987). The living 
hint-fish Ncoceratndus and plotosid callish have 
also been recorded (Kemp & Molnar 1981; 
Mackness et al 1999). Reptile remains described 
so far pertain lo larger ia*a, such as crocodihans 



(Palimnarchus), turtles and madtsoud snakes 
(Bartholomai & Woods 1976. Gal'fney 1981. 
Gaffney & Bartholomai 1979; Mackness & 
Scanlon 1998; Woods I960. 1962). The only 
lizards to date have been Hecht's (1975) 
identification of a number of large varanid fossils 
from the Chinchilla Local Fauna as Megulania sp. 

The Chinchilla Local Fauna appears to 
biocorrclatc with the Kanunka Local Fauna of the 
Tirari Desert, South Australia (Tedford et al 1992) 
and the Spring Park, Bow and Big Sink Local 
Faunas (Mackness et al 2000). On the basis of the 
Kanunka Local Fauna having a 
magnetostratigraphic age of approximately 3.4 
million years, the Chinchilla Local Fauna is 
estimated to be between late Early Pliocene and 
Middle Pliocene in age (Tedford et al 1992). 

Lizard fossil remains have received relatively 
1 ill lc attention in reports on Australian fossil 
faunas, due hoth to a paucity of material and a 
lack of data concerning osteology In some 
lineages of lizards, such as agamids (Covaeevieh 
et al 1990) and varanids (Mackness & Hutchinson 
2000), the restricted amount of morphological 
divergence within the living fauna makes 
identification problematic. For ihe anatomically 
diverse skinks and gckkonoids, this picture is 
steadily changing for the better as new finds focus 
attention on specific areas of anatomy. In this 



170 



MN HUTCHINSON & BS MACKNESS 



study we point out some useful character slates of 
gekkonid skulls that permit attribution to both 
major lineages and more restricted species groups. 
We also build on the knowledge of one of 
Australia's most characteristic lizard faunal 
elements, the bluetongue skinks of the genus 
Tiliqua (Shea & Hutchinson 1992) to describe a 
new and distinctive species. 



Materials and Methods 

The specimens described here were recovered 
through quarrying or wet sieving of sediments 
from "Wilkinson's Quarry', Chinchilla, 
Queensland, a site that has been continuously 
worked by Cec and Doris Wilkinson for over 1 1 
years. The sediments are primarily fluviatile in 
nature and represent a number of depositional 
events. Most fossils in these units occur as 
isolated pieces and all represent Pliocene taxa 
(Mackness, Wilkinson & Wilkinson 1999). 

Specimens were examined using a Wild M3Z 
stereomicroscope with eyepiece micrometer and 
drawing tube. The fossils are registered in the 
collection of the Queensland Museum, Brisbane 
and were compared with skeletal material in the 
collection of the South Australian and Western 
Australian Museums. Comparative scincid 
specimens included all living species of Tiliqua 
and representatives from all species groups in the 
genera Cyclodomorphus and Egernia. The 
morphology of the gekkonid fossils dictated the 
gekkonid specimens that would be the most likely 
candidates for comparison, the 'unspecialised' 
small- to medium-sized diplodactylines. We also 
examined representatives of those genera of 
gekkonines that occur in the Australian region and 
the adjacent Melanesian-Indonesian region, as 
well as a scattering of specimens from across the 
global diversity of the subfamily Gekkoninae. In 
addition, published diagrams and discussions of 
gekkonid cranial osteology were consulted to 
ensure that our selection was representative, 
mainly aimed at justifying some of the general 
statements we make regarding major patterns of 
variation that differentiate gekkonine and 
diplodactyline skulls and mandibles. Specimens 
examined are listed in Appendix 1. Terminology 
for bones follows Estes et al (1988). 

Measurements 

Measurements of varanid vertebrae and the 
large Tiliqua species were made using Vernier 
callipers accurate to 0.05 mm, while smaller 



specimens and teeth were measured with a 
calibrated microscope eyepiece. Vertebral 
measurements are summarised below and largely 
follow Smith (1976). Statistical analyses of these 
measurements are provided in Appendix 2; Tables 
1 and 2. 

Vertebral length (Pr-Po) — the greatest distance 
from the anterior edge of the prezygapophysis to 
the posterior edge of the postzygapophysis. 

Vertebral width (Pr-Pr) — the maximum width 
of the vertebra between the lateral edges of the 
prezygapophyses. 

Vertebral width (Po-Po) — the maximum width 
of the vertebra between the lateral edges of the 
postzygapophyses. 

Centrum minimum width (BW) — the smallest 
distance across the centrum. 

Condylar width (CW) — the greatest transverse 
diameter of the condyle. 

Abbreviation for specimen numbers: AR: 
University of New South Wales Research 
Collection; SAM: South Australian Museum; 
WPC: Wilkinson Private Collection. 



Systematics 

Order SQUAMATA Oppel, 1811 

Family AG AMID AE Gray, 1827 

Unidentified material 

Material examined 

A left dentary (WPC 1354), bearing mid to rear 
section of tooth row. 

Characters 

Agamid reptiles are distinguished in having a 
dentition combining one to three anterior 
pleurodont teeth followed by acrodont teeth. Other 
agamid features are summarised by Estes (1983). 

Description 

The specimen is from a moderately sized 
individual, perhaps of skull length of approximately 
30 mm, and bears nine acrodont teeth. The 
posterior of the dentary is largely missing although 
the facet for the coronoid is still visible. No mental 
foramina are discernible on the specimen. 

Remarks 

Covacevich et al (1990) discussed several 
problems in identifying fragmentary agamid 
remains beyond family level. 




QUEENSLAND PLIOCENE FOSSIL LIZARDS 
pr. dors. 



pr. vom. 



171 



pr. st 




muse. 



fac. cor. 



fac. art. 




fac. cor. 




f. men. 



FIGURE 1. A-F Diplodactylus cf steindachneri. A, eight maxilla (QM F30573) in lateral view and B, medial view_ 
C. left parietal (QM F30574) in dorsal view. D-F right mandible (QM F30572). D, lingual view of dentary. E, 
lingual view of articular + surangular. F, articulated mandible in labial view. Abbreviations: ch. sh. - choanal shelf; 
f. men. - mental foramina; fac. art. - articular facet; fac. cor. - coronoid facet; fac. j. - jugal facet; fac. spl. - 
splenial facet; muse. - insertion point for dorsal neck musculature; pr. ang. - angular process; pr. cor. - coronoid 
process; pr. dors. - dorsal process; pr. st. - supratemporal process; pr. sur. - surangular process; pr. vom. - 
vomerine process; rap - retroarticular process; spl. n. - splenial notch. Scale bar = 5 mm. 



Family GEKKONIDAE Gray, 1825 

Subfamily DIPLODACTYLINAE Underwood, 
1954 

Diplodactylus Gray, 1825 

Diplodactylus cf steindachneri 
(Fig. 1A-F) 



Material examined 

Right maxilla (QM F30573); left parietal (QM 
F30574); partial right mandible (QM F30572), 
consisting of a dentary and fused surangular plus 
articular. 

Description 

Maxilla. The maxilla (QM F30573, Fig. 1A-B), 
a right element, is almost complete, with slight 



172 



MN HUTCHINSON & BS MACKNESS 



erosion of the edges on the margin of the dorsal 
process. Total length 6.1 mm. A slender, 
dorsoventrally flattened vomerine process extends 
medially from the anteromedial end of the bone. 
The facial portion of the maxilla is gently arched 
along its dorsal margin. Although the dorsal 
margin is slightly damaged, there is no 
posterodorsally directed frontal process. The 
margin bordering the nasal opening is almost 
semicircular. The outer surface of the maxilla 
bears two series of foramina; one consisting of a 
row of six openings just above the teeth, the 
posleriormost the largest; and a second consisting 
of three relatively large openings at the level of 
the anteroventral corner of the orbit. The 
subocular ramus of the maxilla is robust and 
relatively short and deep. It does not taper to a 
point, as is usual in gekkonoids; the outline of the 
caudal extremity of the specimen is bifurcated, 
with the dorsal fork larger than the ventral. The 
internal surface of the maxilla bears a prominent 
choanal shelf, which is constricted at its mid-point 
and again anteriorly before projecting as the 
vomerine process. The inner face of the suborbital 
region has a small facet for a vestigial jugal on the 
dorsal fork, below which is a longer facet for the 
ectopterygoid. 

There are 33 tooth loci, most with intact teeth. 
The teeth are small, cylindrical in section and 
tapering to acute points, with an apical cusp and 
lingual cusp separated by a narrow occlusal 
groove; this morphology is regarded as typical and 
plesiomorphic for gekkonids (Sumida & Murphy 
1987). The individual teeth are uniform in size 
and vary only slightly in shape. 

Parietal. The left parietal (QM F30574) (Fig. 
1C) is complete except for slight erosion along 
the medial margin. In common with many living 
species, the medial edge of the bone may not 
have been fully ossified. Beginning at the bone's 
anteromedial apex, the anterior margin describes 
a shallow sinusoidal curve posterolaterally, 
terminating at the obtusely pointed apex of the 
articulation point with the postorbital. The 
caudal margin of the body of the parietal is 
almost straight-edged and faces caudolaterally. 
This margin bears a shallow caudally opening 
pocket, which would have been the insertion 
point for the superficial neck musculature. The 
supratemporal process is relatively slender and 
tapers to an acute point. Ventrally, the bone 
bears a descending flange parallel to the lateral 
margin, the flange descending to a pointed 
projection (epipterygoid process) about halfway 
along. Overall length of the specimen is 4.2 mm; 



width (measured at the apex of the postorbital 
articulation) 1.9 mm. 

Adult mandible (QM F30572, Fig. 1D-F) is 
represented by a complete dentary and nearly 
complete compound bone consisting of the fused 
articular and surangular bones. The angular, 
splenial (or the composite angulosplenial typical 
of diplodactylines) and coronoid are absent. 

Dentary. Total length (taken as a straight line 
from symphysis to tip of angular process) 6.5 mm. 
There are 36 teeth or tooth loci, the teeth being 
similar in size and shape to those on the maxilla, 
with the most anterior teeth having somewhat 
more recurved tips. 

The dental sulcus is well defined by a lingual 
parapet that is evident as far as the posteriormost 
tooth. The posterior limit of the bone has three 
processes, which define two embayments. The 
uppermost (coronoid) process terminates just 
behind the last tooth and has facets on its lingual 
and labial aspects for the coronoid bone. The 
middle (surangular) and lower (angular) processes 
would have terminated at about the level of the 
apex of the dorsal process of the coronoid bone. 
The labial face bears four mental foramina, the 
posteriormost lying at the level of the 24th tooth. 
On the lingual face, the splenial notch extends 
forward to the level of the 27th tooth, and there is 
a facet extending anteriorly to the notch, 
suggesting that the splenial terminated at about 
the level of the 22nd tooth. There is no bony 
intramandibular septum exposed within the 
splenial notch. 

Comparisons 

The identification of these specimens as 
gekkonoids is based on several characters that are 
uniquely combined in this clade. The morphology 
of the articular and retroarticular region of the 
mandible shows an articular facet facing 
posterodorsally and the retroarticular process as 
slender and ventrally positioned. There is 
complete overgrowth by the dentary of the groove 
for Meckel's cartilage. The teeth are small, finely 
pointed and numerous. The parietal is poorly 
ossified, paired and lacks a pineal foramen. The 
elements are those of a small (approx 45 mm 
SVL) generalised gecko. 

Studies of the phylogenetic relationships among 
the Gekkota (Bauer 1990; Donncllan et al 1999; 
Estes et al 1988; Kluge 1967a, 1967b, 1987) agree 
that the Australian region has three of the four 
major gekkotan lineages: the Gekkoninae, 
Diplodactylinae and Pygopodidae. The remaining 
gekkotan clade, the Eublepharidae, is found no 



QUEENSLAND PLIOCENE FOSSIL LIZARDS 



173 



closet than Borneo (the genus Aelurascalabotes). 
To further establish the affinities of the Chinchilla 
Specimens, we have noted several characters that 
appear to be useful in diagnosing membership of 
particular gekkolan lineages, based on the most 
commonly recovered elements: mandibles, 
maxillae, fronlals and parietals. 

Maxilla, The shape of the maxilla differs 
markedly between most gekkonincs and the rest 
of the gekkotan lineages. Authors describing this 
area of the face generally emphasise bony 
coniacts, using the dichotomy of either a 
prcfrontal-nasal contact (shown by most 
diplodactylines) or a frontal-maxilla contact 
(shown by most gekkonines) (Fig. 2). In most 
Australian gekkonines (Cyrtodactylus, Geltyra, 
Heteronotia, Nactus), and in most other 
gekkonines examined, the maxilla arches 
poslcromcdially to contact the frontal, terminating 
in a frontal process that greatly reduces the 
exposure of the prefrontal and excludes the 
prefrontal from the nasal, In the remaining 
Australian gekkonine Christinas and in 



Diplodactyline 



diplodactylines, the maxilla lacks this frontal 
process and the prefrontal includes un 
anteromedial process that contacts the nasal, 
usually excluding it from the maxilla. In 
eublepharids {Aelurascalabotes (Grismer 1088], 
Hemitheconyx [Rieppel 1984]) the prefrontals 
bear anteromedial processes bul they are excluded 
from the maxilla due to well-developed 
anterolateral processes of the frontal; the maxilla 
is similar in shape to thai ot the diplodactylines. 
Pygopods arc variable but most lend to show a 
gekkonine- like posierodorsal expansion of the 
maxilla lo contact the frontal. The maxilla of the 
Chinchilla gecko has a low, gently arched dorsal 
process with no frontal process. 

The Chinchilla fossil shows a distinctive feature 
of the maxilla that suggests a particular 
relationship within the Diplodactylinae. In lateral 
view its subocular ramus shows a relatively deep, 
bifurcate posterior termination, a characteristic 
also seen in a few species of the genus 
Diplodactylus. In these species the notched tip of 
the terminal portion of the maxilla is correlated 



Gekkonine 




FIGURE 2. Comparison of the anterior part ot the skull in a diplodactyline (Diplodavtylus xieiit'chntylus) and an 
Australian gekkonine (Gehyra duhia) gecko showing Ihc usual patterns of contact between Ihe frontal (f), nasal (n), 
maxilla (mx) and prefrontal (pr(') bones. Areas in grey show the anteromedial (nasal) process of the prefrontal 
(diplodactyline), and the frontal process of the maxilla (gekkonine). D. srenodacylus also shows a vestigial jugal 
(j ). compared with the usual degree of development of (his bone in most oiher geckkotans as shown by 6". dubia. 



174 



MN HUTCHINSON & BS MACKNESS 



a 




JJ 5 ~i -> ° * 




T-Trr^rwm^Y^TW~Tw 



B 



?; 




Tnnnrrrvvuwv^ri^ 



— -in? 




— — > j 



• ^ 




FIGURE 3. Right maxilla (slightly anterodorsal in aspect) and sketch of dorsal view of right orbit in three species 
of Diplodactylus, showing reduction of the jugal (grey), A, D. granariensis (R29135); B, D. damaeus (R24553): 
and C, D. steindachneri (R05153). Approx. length of maxillae 6-7 mm, not to scale. 



with reduction of the jugal bone reported by 
Kluge (1967b). The two species reported as 
having the greatest reduction of the jugal (D. 
steindachneri and D. stenodactylus) have the most 
pronounced development of the bifurcate shape 
(Fig. 3A-C) and are essentially identical in 
appearance to the Chinchilla maxilla. 

Frontal. No frontal has been recovered at 
Chinchilla, but future finds should be identifiable 
as gekkonine if there are three facets on its 
anterior portion (for the nasal, maxilla and 
prefrontal) or otherwise if there are only two 
facets. The relatively long nasal processes of 
eublepharid frontals might also be diagnostic for 
that clade. 

Parietal. Many gekkotan taxa have distinctive 
parietal morphologies. However, this bone shows 
marked ontogenetic variation in its degree of 
ossification, the elaboration of muscle attachment 
points on its lateral and posterior margins, and the 
robustness of the supratemporal process. The most 
distinctive feature of the Chinchilla specimen is 
the concavity for the neck musculature on the 
posterior margin. This arrangement is not typical 



of most geckos, where the musculature simply 
attaches to the posterodorsal surface and edge of 
the parietal, or to the posteroventrally deflected 
trailing edge, generally with no more than a 
depression or a low ridge to mark the point of 
attachment. A more-or-less well-defined pocket is 
present in some members of the genus 
Diplodactylus (D. stenodactylus, D. steindachneri, 
D. byrnei, intraspecifically variable in D. 
damaeus) but is absent from other Diplodactylinae 
examined and was not seen in the gekkonines. 
Pygopod parietals are much slenderer than those 
of other gekkotans, the midline length being at 
least twice the width (cf less than 1.5 times the 
width). 

Mandible. Some diplodactyline and gekkonine 
taxa as well as pygopods (Hutchinson 1997) have 
unique modifications of the mandible that are 
diagnostic for individual genera, but many 
gekkotan taxa have mandibles that are 
superficially similar in overall shape. In these 
generalised mandibles, the retroarticular process 
provides a key to the two major gecko 
subfamilies. In gekkonines, the retroarticular 



QUEENSLAND PLIOCENE FOSSIL LIZARDS 



175 



process is generally spoon-like in shape, its dorsal 
surface forming an obvious, concave bowl. In 
most diplodactylines (Sultuarius and some 
Strophanti are exceptions) and in pygopodids, the 
retroarticular process is rod-like, with an oval, 
flattened or shallowly concave cross-section and 
terminated by a club. The morphology of the 
dorsal surface of the retroarticular process in 
eublepharids has not been described (published 
diagrams show only the shape in lateral or ventraJ 
view). The Chinchilla specimen has the clubbed, 
diplodactyline type of retroarticular process. 

The fossil maxilla and mandible therefore not 
only show an overall similarity with 
diplodactylines, but also share several discrete 
character states that indicate specific 
diplodactyline affinities. The characteristics that 
distinguish the fossils are those that are uniquely 
combined in the genus Diplodactylus. 

The Chinchilla fossils were found as 
disarticulated elements but their size and 
preservation, especially the mandible and maxilla, 
are consistent with having come from one animal. 
Based on the possession of a bifurcate terminus to 



the posterior ramus of the maxilla and the 
presence of a 'pocket' on the trailing edge of the 
parietal, they are referred to D. steiiuiachneri. It is 
one of three species of Diplodactylus still found 
in the region (the other two are D. xittatus and D. 
tessellaius, Ingram & Raven 1991), and is the only 
one of these three species to have a vestigial jugal 
and bifurcate maxillary terminus. The other 
species with reduced jugals, members of the D, 
stenodactylus group, are found in the deserts of 
central and western Australia, the closest to 
Chinchilla being D. immaculatus of western 
Queensland. 



Family SCINCIDAE Gray, 1825 

Subfamily LYGOSOMINAE Mittleman, 1952 



Tiliqua Gray. 1825 

Tiliqua wilkinsanurum sp. nov. 
(Fig. 4A-B) 



pr cor 



pr. ang 



sym 




f. men. 



s. sur. 



cr. sym 



can. Inf. alv. 



spl. n. 



FIGURE 4 A-B Tiliqua wilkinxonorum n. sp. QM F30567 Holotypc right dentary. A, labial; B, lingual. 
Abbreviations; can. inf. alv. ~ inferior alveolar canal; cr sym. - symphysial crest; f. men. - mental foramina; fac. 
Cor. - comnoid facet; iaT — inferior alveolar foramen (= anterior inferior alveolar foramen); mis - inlramandibulur 
septum; mames - limits of attachment for external adductor musculature; pr. ang. - angular process; pr. cor. - 
coronoid process; spl. n. - splenial notch; s. sur. - surangular suture, sym - symphysis. Scale bar= 5 mm 



176 



MN HUTCHINSON & BS MACKNESS 



Material examined 
Holotype: a right dentary (QM F30567). 

Type Locality 

Wilkinson's Quarry site, Chinchilla area, 
southeastern Queensland. The specimen comes 
from a fossil-bearing unit within the quarry that 
lies unconformably on top of a layer of fine sand. 
The sediments are primarily fluviatile in nature 
and represent a number of depositional events. 

Age 

Chinchilla Local Fauna, late Early to Middle 
Pliocene (Tedford et al 1992). 

Diagnosis 

A large species, dentary tooth row length 
33 mm, differing from other Tiliqua by the 
following combination of characters: all tooth 
crowns pointed and retaining an occlusal ridge, 
the crown scarcely wider than the shaft; coronoid 
process of the dentary relatively slender, narrower 
than long, its shape correlated with a curved and 
concave, rather than angular or straight, dentary- 
surangular suture connecting the base of the 
coronoid process with the tip of the angular 
process. 

Description 

A right dentary, bearing a complete row of 20 
teeth or tooth loci. The specimen is almost intact, 
being slightly damaged at the apex of the splenial 
notch and on the ventral edge adjacent to a 
repaired crack through the jaw at the level of the 
15th tooth. Total length, from the anteriormost 
point of the symphysis to the tip of the angular 
process, 47.8 mm. Depth al level of 16th tooth 
(excluding tooth), 7.8 mm. Maximum depth of 
jaw, measured vertically from the apex of the 
coronoid process, 13.1 mm. Maximum width, 
measured in occlusal view at the level of the 1 6th 
tooth, 8.0 mm. 

Meckelian groove closed, no trace of suture. A 
pronounced crest runs caudally from the 
symphysis along the ventrolingual margin of the 
jaw, gradually merging with the body of the 
dentary at about the level of the eighth tooth. 
Apex of the splenial notch at about the level of 
the 12th or 13th tooth, slight uncertainty due to 
the broken edge of the inferior alveolar foramen 
at the apex of the notch. Labial surface with 
pronounced, arcuate adductor muscle scar that has 
its apex level with the last tooth. Angular process 
projects 15 mm beyond the level of the last tooth. 
Dorsal margin of angular process merges with the 



ventral margin of the coronoid process along a 
smooth concave curve that has its apex at the level 
of the tooth row. Coronoid process well 
developed, flattened, but relatively small 
compared with other Tiliqua. A series of mental 
foramina starts at the level of the 1 1th tooth and 
runs anteriorly to the symphysis. These foramina 
are mostly arranged as a series of superposed 
pairs, with a total of about nine in the series. 

Intramandibular septum fused ventrally along 
its length to the ventral lamina of the dentary, 
completely separating the inferior alveolar canal 
from the Meckelian canal. Caudal margin of 
intramandibular septum with a deep, curved 
notch, bounded ventrally by a caudally projecting 
prong. 

Tooth row, in occlusal view, almost straight, 
gently curving medially anterior to the level of the 
sixth tooth. Labial wall of dental sulcus tall and 
robust. Lingual wall low but distinct anteriorly, 
reducing caudally and disappearing by about the 
level of the 16th tooth. Lingual face of dentary 
below the dental sulcus vertical, sharply distinct 
from the tooth row. Dentition pleurodont. The 
first of 20 teeth is broken off and the 10th, 12th 
and 19th loci are empty, but all other teeth are 
present and well preserved. The teeth vary 
distinctly in size, becoming larger progressing 
caudally, with the maximum reached in positions 
13 to 16, the subsequent teeth then diminishing in 
size. As a guide to the changes in tooth size, tooth 
6 is 3.5 mm high by 1.3 mm wide (measured 
normal to the lingual aspect) while tooth 14 is 
4.3 mm high by 2.4 mm wide; thus, height 
increases by 23% and width by 85%. In lingual 
view, the tooth crowns are acutely pointed 
anteriorly, becoming obtusely pointed by about 
the 13th tooth. The crown of each tooth is 
margined by an occlusal ridge that separates its 
lingual and labial surfaces. In mesial view the 
tooth crowns have a parabolic curved outline 
labially, but the lingual surface is flatter, 
descending almost vertically from the apical ridge 
and forming an angular contact with the rounded 
labial surface. On the largest teeth, several low 
striae run vertically down from the apical ridge on 
both labial and lingual surfaces of the crown. 

Comparisons 

The combination of the closed Meckelian 
groove, enlarged 'cheek' teeth with more than one 
large tooth posterior to position 10, and an 
enlarged and flattened coronoid process is unique 
and diagnostic for the bluetongue lizards of the 
genus Tiliqua (Shea 1990). The related genus 



QUEENSLAND PLIOCENE FOSSIL LIZARDS 



m 



Cyilvdonwrphns has only a single enlarged cheek 
toothy while other lygosornines wnh a closed 
Mcckelian groove Jack such obvious denial 
differentiation. 

Most species of Tiliqua, however, have a more 
specialised dentition than T. wdkinsonorum. The 
plesiomorphic loolft crown morphology m skmks 
includes a convex outer surface, curving 
occlusal ty to a pair of apical ridges separated by a 
grndVB (M. H. pers. nbs.; Sunnda & Murphy 
1987), while the lingual surface is flattened and 
oriented almost vertically, forjiung an angular 
contact with Ihe apical ridge, In lingual and mesial 
views lite crown is acutely pointed. Jn contrast to 
Ihis. species of CyctcuUimarphus and Tiliqua have 
teeth modified For dumphagy (Estes &- Williams 
1984). Most species of Tihqua, including T, 
giga*. T. multifasciala, T occipitalis, T. pusilla, 
] n.tqosa arid "/. scmcoides, have cheek teeth with 
expanded crowns, and have lost or greatly reduced 
the plesiotnorpriic asymmetry of ihe crown. In 
occlusal view the crown is circular in shape, and 
when viewed mesicdistaJly the crown profile, a 
convex curve is sitndar in shape ungual!) and 
lahially. The apex ai the rrmfh has a central 
conical projection with pronounced slnae 
tadiaung over liic crown from this central poim. 
Most individuals lack an occlusal ridge, although 
many T occipitalis ami some T rugosa have a 
remnant on the tooth apex. Tiliqua adelaidensis 
has tjuite different dentition in which lite enlarged 
cheek teeth are labiolingually ce-mpresscd, with a 
pronounced cutting edge formed by the apical 
ridge. Tiliqua nigrolutea is the living species thai 
shows the- closest dental similarity to T 
wdhnsonorurn. The tcclh show lutle expansion of 
the crowns, which are pointed a/id retain an 
obvious occlusal ridge that demarcates the crown 
into lingual and labial aspects. These two species 
show an asymmetric tooth crown profile in 
inesiodisul view, with a more steeply descending 
lingual surface. Compared with T. ni±:r.->lutca. the 
teeth of T. wdkinsonorum differ only in being 
more robust and having ihe crowns of the largest 
leeih more expanded and obtusely pointed. These 
dilleccnecs could be attributed to allometry nr 
individual variation in tooih ppjportions, which is 
considerable in living Tiliqua (M. H. pers. obs.l. 
However, other characteristics of ihe fossil 
.specimen differentiate it from both T tngrvhttvu 
and other Tdtqiui species. 

The dentary is from a very large Tiliqua L r sing 
llic average ratio of tooth row to mandible length 
(0.48) that applies in extant Tiliqua. Ihe loolh row 
length of 7. wdkinsonorum of 33 mm extrapolates 



to an approximate mandible length of 68 nun. 
This si«r is approached only by the largest males 
of T. rugosa and T. scincoides (G. M. Shea pers. 
eomm. ) and suggests that T. wdkinsonorum may 
have had a snout-vent length close to 400 mm and 
a mass in excess of one kilogram. In spite of this 
size, the dentary of T. wilkinsanorum is noiably 
gracile. This is most clearly seen in the shape of 
the coronoid process and the surangular suture 
running from the base of the coronoid proce (5 10 
the lip of the angular process. In all exlanl Tiliqtta, 
the coronoid process is dorsally and caudally 
expanded This expansion is masked to some 
cxtenl because the base of the coronoid process is 
absorbed by caudal expansion of Ihe trailing edge 
of the dentary such thar the surangular suture is 
not deeply incised into the labial lamina of Ihe 
dentary. The precise course of this suture varies 
both inter- and intraspecillcaily (Fig. 5A-1). In T 
nigrotuiea, T. occipitalis and T. rugosa, this 
suture usually runs vertically from ihe coronoid 
process and then angles caudoventrally to the tip 
of the angular process of the dentary. In 7. 
adelaidensis, T gigas. T. mulitfasciutu and 71 
xcinoiiJi'-;, 'he suture fends lo run caudovenlrally 
in an almost straight line from the base ot Che 
coronoid process. In none does it trace ihe 
excavated curve seen in T. wdkinsonorum fund 
olher skinks). In T. wdkinsonorum the coronoid 
process is less expanded than in any exlanl 
species, with the possible exception of T. gtgat, 
but is more clearly defined ventrally by the 
anterior sweep of the concave surangular suture. 
In living Tiliqm species, the expanded coronoid 
process and adjacent parts of the denlary increase 
ihe strength of this region of the mandible and 
increase the area of attachment for the jaw-eJosing 
tendon of the m. adductor medius extemus 
uqitrficialis (Haas 1973). The relative slendcracss 
of the coronoid pioeess and ihe concave 
surangular sulun* in if! wdkinsonorum are C 
to the morphology seen in oilier skmks, including 
Egernia. the genus that probably includes the 
sister group of the Tiliqua lineage. 

In summary, T. wdkinsonorum is one of the 
largest known specimens of Tiliqua, and has less 
Specialised tcclh than all but T. nigrolutea and a 
more slender (plesiomorphic) coronoid process of 
the dentary than all but T. gigus. The morphology 
of the teeth and the relative gracdity of the caudal 
end of the dentary suggest that 7'. wdkinsonorum 
was not as specialised for duroplmgy as most 
other large Tiliqua. 

Tiliqua wilkinsonorum is the third species of 
Tiliqua lo be found in the Tertiary of Australia 




MN HUTCHINSON & BS MACKNESS 



B 










FIGURE 5. Dentary variation in Tiliqua. Detailed views showing variation evident in the degree of expansion of the 
coronoid process of the dentary and the shape of the dentary-surangular suture. All to same scale except T. 
adelaidensis (inset); both scale bars = 5 mm. A, T. wilkinsonorum. B, T. rugosa R27028. C, T. adelaidensis 
R40738. D, T. nigrolutea R02725. E, T. nigrolutea R47698. F, T. occipitalis R35758. G, T. scincoides R27039. H, 
T. gigas R11419. I, T. multifasciata, R35757. 



and the second extinct species. Shea and 
Hutchinson (1992) described a very small species, 
T. pusilla, from the early Middle Miocene of 
Queensland, and specimens indistinguishable 
from a living species, T. scincoides, have been 
recorded from the Pliocene of South Australia 
(Pledge 1992) and Queensland (Mackness & 
Hutchinson 2000). 



cf. Cyclodomorphus Fitzinger, 1843 

Material examined 

A partial dentary (QM F30568). 

Description 

The specimen represents about one-third of a 
left dentary, lacking the anterior portion, anterior 
to the last six teeth and lacking the coronoid and 
angular processes. It is not a juvenile, based on 



several cycles of tooth replacement being evident. 
The groove for the Meckelian cartilage is 
completely obliterated by the dentary. The third- 
last tooth is markedly enlarged. The tooth anterior 
to this is absent, but its locus is smaller, while the 
other remaining teeth are smaller again. A single 
large mental foramen is present on the labial 
surface of the jaw at the level of the fourth-last 
tooth. The teeth, both enlarged and small, have 
blunt, slightly laterally compressed crowns with 
virtually no apical ornamentation such as cusps, 
grooves or striae. 

Comparison 

The fossil appears to have only one markedly 
enlarged cheek tooth, with a second moderately 
enlarged tooth anterior to it. Among extant 
Australian skinks, only members of the Tiliqua 
lineage combine a closed Meckelian groove and 
enlarged durophagous cheek teeth (Shea 1990). 



QUEENSLAND PLIOCENE FOSSIL LIZARDS 



179 



The dentitions of the two genera of this lineage, 
Tiliqua and Cyclodomorphus, differ in that adults 
of Tiliqua have several enlarged cheek teeth while 
those of Cxclodo/narphus usually have only one 
strongly enlarged tooth. On this basis as well as 
its small size and overall shape, the Chinchilla 
specimen is more similar to Cyclodomorphus than 
it is to Tiliqua. However, Tiliqua adetaidensis and 
the extinct T. pusilla show that the small species 
of Tiliqua may have only two enlarged cheek 
teeth, so that the distinction on tooth pattern alone 
is not entirely conclusive. Other attributes 
distinguishing the two genera, including the shape 
of the symphysial region, are not: preserved on the 
specimen. In the shape of the crowns and lack o( 
occlusal striae, the fossil js unlike any species of 
either Cyclodomorphus Or Tiliqua examined in 
this sludy, but it is too fragmentary to either 
allocate to a genus with certainty or to warrant 
description as a new faxon. 



Family VARAN1DAE Cray. IS27 

IMegalania sp. 

Material rxamincil 

Five isolated dorsal vertebrae (WPC 115, 116, 
2409. 2014, 3322), three isolated dorsal fragments 
(WPC 2040, .3555, 3556), four isolated caudal 
vertebrae (WPC, 1429, 1573, 2364, 2524). 

Clio ratters 

The genus Megalania is characterised in part 
by having massive thoracic and lumbar 
vertebrae with weakly developed zygosphenes 
(absent in typical Varanus) as well as small 
depressed neural canals. The adult teeth of 
Megalania are large and slightly recurved 
distally. The anterior cutting edge is rounded 
and serrated distally. The posterior cutting edge 
is thin, blade-like and serrated along its entire 
length (Hecht 1975) 

Description 

The dorsal vertebrae are much more massive 
than those of any extant varanid but compare well 
in morphology with those from an extant varanid 
Varunus sarins (AR 7641) as well as from large 
fossil varanids from the Bluff Downs Local Fauna 
(Appendix 2: Tables 1 and 2). The Chinchilla 
specimens are 62% larger than these 
measurements supplied lor Australia's largest 
extant varanid, the perentie Varanus giganteus by 
Smith (1976) and 20% larger than those for I he 



large fossil varanid from Blufr Downs Local 
Fauna (Mackncss & Hutchinson 2000). 

Remarks 

A single maxilla (QM F874) from the 
Chinchilla Sand was referred to Varanus dirus 
(Hecht 1975), a taxon originally described by De 
Vis (1889) on the basis of an isolated tooth from 
the Pleistocene Kings Creek locality. Hecht (1975) 
synonymised V. dints with Megalania prisux 
Four isolated vertebrae (two caudals and two 
dorsals) have also been collected from the 
Chinchilla Sand and, although Hecht (1975) 
referred them to Megalania, he also suggested that 
this material may represent a separate Pliocene 
species. 

The assignment of the larger varanid vertebrae 
described in this paper to IMegalania is done 
purely on the basis of convention. Opinion is 
divided as to the current status of the genus, with 
Hecht (1975) and Molnar (1990) concluding it is 
valid but Estes (1983) and Lee (1996) suggesting 
ihat Megalania should be synonymised with 
Varanus. A large collection of varanid fossils 
from Pliocene localities is presently under study 
by the authors. 



Vatanus sp. 

Material examined 

Two isolated dorsal vertebrae (WPC 118, 
1430), one isolated caudal vertebra (WPC 3557). 

Characters 

These vertebrae are identified as varanid by 
their distinctive condyle-cotyle articulations. The 
dorsal vertebrae also show the characteristic 
constriction of the centra anterior to the condyle 
seen in Varanus. 

Description 

The vertebrae show characteristic varanid 
motphology with a range of centra lengths 
(Appendix 2; Table 3) that indicate a medium- 
sized goanna of the size of Varanus gauldii 

Remarks 

Although Wilkinson (1995) suggests some 
characters for separating varanid species on the 
basis of vertebral morphology, many of these rely 
on the neural spine being present. This feature is 
missing on all three fossil vertebrae. Interspecific 
and/or intraspecific variation in varanid vertebrae 
have yet to be fully tested, iheteforc, characters 



180 



MN HUTCHINSON & BS MACKNESS 



identified by Wilkinson (1995) must be used with 
caution. 



Discussion 

The composition of the lizard fauna from the 
Chinchilla Sand Local Fauna, at least at family 
level (Appendix 2; Table 4), is similar to that of 
the Bluff Downs Local Fauna. This is the only 
other Australian Pliocene locality with a 
significant lizard record (Mackness & Hutchinson 
2000). It differs from Bluff Downs in the generic 
representatives of each family, but the significance 
of this is unknown; it could merely be stochastic, 
reflecting the very small sample sizes at each 
locality. The palaeoenvironment was primarily 
fluviatile for Chinchilla and lacustrine and 
fluviatile for Bluff Downs. 

The exact nature of the Chinchilla 
palaeoecology has yet to be determined. While 
there may have been a significant wetland 
component, as suggested by the presence of 
various aquatic taxa such as turtles (Gaffney 1981; 
Gaffney & Bartholomai 1979) and waterbirds 
(Olson 1975, 1977), there is also evidence that the 
environment was highly seasonal (Mackness et al 
1999; Wroe & Mackness 2000). The presence of 
arboreal species such as Koobor (Archer 1977) 
and an unnamed phascolarctomorph (Mackness et 
al 1994) would be consistent with a complex and 
mature forest, but the composition of the 
mammalian fauna indicates a lack of undoubted 



closed-forest species. Further, the predominance 
of grazing kangaroos and diprotodontids suggests 
that grasslands were also part of the environment. 
The presence of an extinct reptile species at 
Chinchilla is noteworthy given the slow rate of 
faunal turnover in reptile populations (La Duke 
1991). Tiliqua wilkinsonorum represents a 
distinctive but extinct species of lizard, a 
relatively uncommon observation for the Pliocene 
(Estes 1983). A conclusion that can be drawn 
from the presence of such 'fossil endemics' is that 
they signal ecological or geographical factors at 
Chinchilla during the Pliocene that are no longer 
present in recent environments. 



Acknowledgments 

The authors thank Cec and Doris Wilkinson for their 
tireless efforts in recovering fossil remains from the 
Chinchilla Local Fauna over the past 12 years and for 
making the lizard material available for us to study. 
Glenn Shea, Jim Mead, Michael Archer and Suzanne 
Hand provided helpful comments on the manuscript. 
The study of the Chinchilla material was supported in 
part by an ARC Program Grant to M Archer; a grant 
from the Department of Arts, Sport, the Environment, 
Tourism and Territories to M Archer, S Hand and 
H Godthelp; a grant from the National Estate Program 
Grants Scheme to M Archer and A Bartholomai; and 
grants in aid to the Riversleigh Research Project from 
the University of New South Wales, Wang Australia 
Pty Ltd, IC1 Australia and the Australian Geographic 
Society. 



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phylogenetic implications. Memoirs of the 
Queensland Museum 45: 641-645. 

Wroe, S & Mackness, BS. 2000b. A new genus and 
species of dasyurid from the Pliocene Chinchilla 
Local Fauna of southeastern Queensland. Alcheringa 
24: 319-326. 



QUEENSLAND PLIOCENE FOSSIL LIZARDS 



I S3 



APPENDIX I 

Ostcological material used for comparisons. Specimens are from the Herpetology Section of the 
South Australian Museum and the Western Australian Museum (WAM prefix). 



Gekkonirtes 

Christinus marmoratus (R35562; R39934), 
Cosymbotus platyurus (R36789), Cyrtodactylus 
louisiudensis (R14002), Gehvra ausiralis 
(R34I36). G. dubia (R42872), G sp. (2n = 44) 
(R44595), G. catenate (R35561), G. oceanica 
(R08687), G. purpurascens (R36377), G, robusta 
(R34222). Gekko gecko (R35559), G japonicus 
(R07435), G. vittatus (R55978) Gonatodes 
humeralis (R40088). Gymnodactvtus darwinii 
(R40090), Hemiductylus frenaius (R35564), 
Ihttraiwtia binoei (R0975I A. R33565, 
R39935), Nactus iheveni (R09740 B). Paroedura 
pirta (R55105), Phelsuma mudaguscartensis 
(R40025), Ptyndactylus hasselquisti (R49804), 
Tarcntola annularis (R40024), Thecuductylus 
rapicauda (R40099-I00). 



Diplodactylines 

ttavayia ornata (R06752), Crenadactylns 
ncellatus (R031I3 C), Diploductyhts byrnei 
(RI35I4), n. conspicillatus (RI9967), D. 
domains (R200I7, R24553) D •granariensis* 
(SA populations) (R020IQ, R26494, R29I35) .') 
immaadatus (R42676), U. ptdcher (R26383), D. 
steindachneri (R05153, R52746), D. 
stenodactylus (R07592, R21240, R26777). D 
tessellatus (R03876, R40938), D. villains 
(R35568), Hopludaciylus pacificus (Rl 1025), 
Nephurus asper (R35567), A', laeiissimus 
(R 14987 A), N levis (ROO309 A, R27044-45), N. 
mitit (R55422, R57083). N. stellalus (R32297), 
Otdura lesueurii (R33597), O. marmorata 
(R4 2893), O. monilis (R35563), O, tnoni 
(R33583), Phyllurus platurus (R35054), 
Rhyruhoedura ornata (R35827), Saltuarius 
swaini (R29205), Strophurus ciliaris (R 10697, 
R35566), 5. intermedins (R 14325 C). 

A range of pygopod skulls was also examined 
(see Hutchinson 1997 for listing). 



Egernia group sktnks 

Carucia zebrata (R35765), Cxclodomorpluts 
nmximus (WAM R77193, WAM R 77637), C. 
michaeti (R35682), C. gerrurdii (R35761, 
R47699), C. melanops elongatus (R03231, 
R03856G, R356S1), C, venustus (R38021), 
Egernia enventryi (R35686, R47693-94), E. 
cuttninghami (R35680, R35763), E. depressa 
(R03433 H), E. formom (WAM R65803), B. 
hosmen (R22510), E. inornate (R07234, R07238, 
R07245A. R35687), E. kingii (WAM R36376. 
WAM R89269-70), E. lucmosa (WAM R36019), 
E. major (R27043, R35762), E mulliscutatu 
(R08469, R25245), E. napoleonis (R35692. 
R51076, WAM R45350), £. pilbarensis (WAM 
R78945), E. pidchra (R40026, WAM R718S4), £. 
richardi (R 10841), E. saxatilis intermedia 
(R43961), E. stokesii (R02560, R41913), E. 
striata (R07I79, R07198), £. striotata 
(R02909B, R 16593, R2689I, R38019). E. whuii 
(R02726, R35690, R27042, R35689, R45316, 
R34886, R35688. R35691), Tiliqua adelaidensis 
(R40738, R40745, R43412), T. gigas (R I 1419), 
r. nudtifasciata (R27041. R35757), T. nigrolutea 
(R02725, R27048, R27050, R47698), T 
occipitalis (R02724. R25369. R27047, R35758), 
T. rugosa (R02563[2J, R02564, R27026, R2702S, 
R02801, R25615, R27027, R27029, R275S4, 
R3I855. R35760), T, scincoides (R02561, 
R27036-40, R35759, R38O20, R43962, R27094). 



184 



MN HUTCHINSON & BS MACKNESS 



APPENDIX 2 

TABLE 1. Measurements (mm) of IMegalania dorsal vertebrae from Bluff Downs and Chinchilla Local Faunas 
compared with Varanus giganteus. Measurements as defined in Methods. Range (mean ± standard deviation). Data 
for V. giganteus taken from Smith (1976), for Bluff Downs from Mackness & Hutchinson (2000). 



Specimen 



No 



Pr-Po 



BW/Pr-Po 



CW/Pr-Po 



Pr-Pr/Pr-Po 



V. giganteus 
Bluff Downs 
Chinchilla 



20 24.5-27.1 (25.7+0.14) 
7 24.5-27.1 (25.7±0.14) 
4 24.5-27.1 (25.7+0.14) 



0.54-0.64 (O.58±.O05) 
0.54-0.64 (0.58+.005) 
0.54-0.64 (0.58±.005) 



0.55-0.63 (0.59+006) 
0.55-0.63 (0.59+006) 
0.55-0.63 (0.59+006) 



0.88-1. 01(0.92±.007) 
0.88-1 .01(0.92±.007) 
0.88-1 .01(0.92±.007) 



TABLE 2. Measurements (mm) of individual IMegalania vertebrae. Measurements as defined in Methods. 



Specimen 



Pr-Po 



Pr-Pr 



Po-Po 



BW 



CW 



COW 



CEL 



WPC116 


39.4 


33.2 


39.3 


20.5 


26.1 


27.6 


25.0 


WPC2014 


38.9 


46.4 


41.4 


- 


- 


30.4 


- 


WPC2409 


48.3 


57.3 


49.5 


26.4 


29.4 


31.6 


28.2 


WPC3322 


37.2 


43.5 


40.4 


17.6 


23.2 


25.2 


26.4 


WPC3555 


56.8 


- 


- 


- 


- 


- 


- 


WPC3556 


44.3 


- 


- 


- 


- 


- 


- 



(Mean/SD) (41.6 ± 6.6) (47.6 ± 6.6) 42.6 ± 4.6) (21.5 + 4.5) (26.2 ±3.1) (28.7 ± 2.9) (26.5 ±1.6) 

TABLE 3. Measurements (mm) of centra of dorsal vertebra of fossil Varanus sp. from Chinchilla Local Fauna 
(WPC), Bluff Downs Local Fauna (QMF) and extant Varanus varlus (AR7641). 



Specimen 


Measurement 


Specimen 


Measurement 


WPC118 


12.2 


AR7641 (a) 


15.3 


WPC 1430 


16.5 


AR7641 (b) 


15.3 


QM F7774 


14.2 


AR7641 (c) 


15.4 


QM F23238 


10.5 


AR7641 (d) 


15.4 


QM F23659 


10.8 


AR7641 (e) 


15.6 


QM F23683 


13.8 


AR7641 (f) 


15.7 



TABLE 4. Comparison of lizard taxa recovered from the Queensland Pliocene (Chinchilla and Bluff Downs Local 
Faunas. Source: Mackness & Hutchinson 2000). Code: present •, absent x, endemic form <e) . 



Taxa 


Chinchilla 


Bluff Downs 


Gekkonidae 






cf Heteronotla 


X 


• 


Dlplodactylus 


• <«> 


X 


Agamidae 






unidentified material 




• 


Varanidae 






IMegalania 




• 


Varanus 




• 


Scincidae 






Tiliqua 


. w 


• 


Cyclodomorphus 




X 


Egernia 


X 


• 


Eulamprus 


X 


• 



FIRST REPORT OF THE CUPRESSACEAN CONES IN THE EOCENE 
EYRE FORMATION OF SOUTH AUSTRALIA 

N, S. Pledge 



Summary 

Conifer impressions are a very minor component in the various 'silcrete flora' assemblages found in 
the interior of South Australia. Previously, no fruiting bodies had been found, although numerous 
foliage species have been recognised. This note describes impressions of the first woody ovulate 
cones found, discovered in a small block of silicified sandstone near Lake Hart, west of Woomera. 
They are distinctive and do not appear to bear close affinity to any modern Australian species, but 
resemble cones of the African Widdringtonia. 



FIRST REPORT OF CUPRESSACEAN CONES IN THE EOCENE 
EYRE FORMATION OF SOUTH AUSTRALIA 



NS PLEDGE 



PLEDGE. NS. 2002. First report of cuprcssaccan cones in llie Eocene Eyre Formation of South 
Australia Records of'ihe South Australian Museum 35(2): 185-191 

Conifer impressions are a very minor component in the various "silerete flora' assemblages 
found in Ihe interior of South Australia Previously, no fruiting bodies had been found, although 
numerous foliage species had been recognised. This note describes impressions of Ihe first 
woody ovulale cones found, discovered in a small block of silicificd sandstone near Lake Hart, 
west of Woomera. They are liislinclive and do nol appear to bear close affinity lo any modern 
Australian species, but resemble cones of the African WiddrinRtania. 

NS Pledge, South Australian Museum, North Terrace, Adelaide, South Australia 50(H). 
Manuscript received 19 April 2002. 



Besides Athrotaxis, the modern Australian flora 
contains three genera of cupressacean conifers: 
Callitris, Aciinostrahus and Diselma. Receni 
morphological and molecular work by Gadek et al 
(2000) shows thai Ihe latter are closely related lo 
an African genus Widdringtonia, within a 
monophyletic group, the subfamily Callitroideae, 
comprising all Southern Hemisphere 
cupressaeeans except Athiota.m. 

Fossil conifers, including members of the 
Cupressaccae, are sparse in the Australian fossil 
record I Hill & Carpenter 1989) Many reported 
specimens have occurred as impressions in fine- 
grained sediments, with consequent difficulty in 
interpreting fine structure, but organic remains 
also occur, eg in Tasmania, and have yielded 
valuable material (ibid). This has more than 
doubled the number of genera in the Australasian 
region, with taxa that occur today in New Guinea, 
New Zealand and elsewhere, as well as several 
extinct genera (Hill & Brodribb 1999). 

Plant impressions in silicificd sandstone have 
been known from the Woomera area / Arcoona 
Plateau for many years (Chapman 1937), and as- 
far as Stuart Creek to the north and Clayton 
Station on the Birdsvillc Track, to the northeast. 
Most of the moulds are of angiosperm leave. 1 . 
(Rowett 1997). a few of which have been 
described (Chapman 1937; Greenwood ct al 
2001). Conifer vegetative shools are a minor 
unobtrusive component, studied by Nunn (1964) 
and Offler (1969, 1984). In the 1970s localities 
were found yielding natural moulds of fruiting 
bodies, mostly of myrtaceous origin (Ambrose et 



al 1979; Lange 1978). A few specimens referable 
to fruit of Proteaceae ifianksia sp.) and 
Casuarinaceae have been noted (Greenwood el al 
2001 ; this writer, unpubl.) but, until now, no 
coniferous cones have been recognised, despite 
the widespread occurrence of vegetative shoots. 
Nunn (1964) recognised 21 different conifer 
morphotypes, some of which could be related to 
modern species from the Auslralasian region. She 
stressed the problems and uncertainties of 
identifying the species 

These plant fossil impressions occur in a 
number of isolated, discrete localities over an area 
of thousands of square kilometres, often in 
apparent channel deposits, eg at Nurrungar, Island 
Lagoon, near Woomera (pers. obs. 1969), m what 
has been regarded as Eyre Formation s.l. Their 
age has long been in dispute (Ambrose et al 1979; 
Callen & Lange 1986; Chapman 1937. 
Greenwood et al 1990: Wopfner et al 1974) but 
recent discoveries at Nelly Creek, Lake Eyre 
South, have at last related them to deposits datable 
by palynological studies to Ihe middle Eocene 
(Alley ct af 1996; Callen & Cowley 1995; 
Chnstophel et al 1992) within the Eyre Formation. 
Variations in the contained fossil flora) 
assemblages suggest that the different localities 
probably have slightly different ages. 



Material and Methods 

In 1979 members of the Woomera Natural 
History Society discovered plant impressions near 



186 



NS PLEDGE 



N 




L. Hart o 



L. for re ns 

^ \ 

Woo mer a 



d 



AUSTRALIA 




dela ide 



FIGURE 1. Locality map, Lake Hart and other 'silcrete 
flora' localities in South Australia. 



Lake Hart in the Woomera Prohibited Area (Fig. 
1). Subsequently, L. Marsh and T. Nurenberg 
presented a slab of quartzite to the South 
Australian Museum (SAM P22732; Fig. 2) 
bearing the impressions of a few leaves and seven 
fruit-like bodies. Silicone rubber casts (Fig. 3) 
taken from these moulds disclosed fruits strongly 
reminiscent of Callitris ovulate cones. Closer 
examination, however, showed that, instead of the 
six subequal valve scales (2 triplets) seen in 
Callitris, these fruit had two unequal pairs of 
valves (Fig. 4). Re-examination of the collection 
of silcrete flora material in the Museum failed to 
uncover any further specimens; SAM P22732 is 
therefore unique. 

Comparison was made with dried specimens in 
the South Australian State Herbarium and with 
trees growing at the Mount Lofty Botanic 
Gardens. 



Description 

Order CONIFERALES 

Family CUPRESSACEAE Neger 

Genus Incertae sedis 

The cones are globose to ellipsoidal in shape, 
with a slightly conical distal apex. The ellipsoidal 
shape may be due to slight compression during 
burial, but the flattening is always the same 
relative to the smaller valve scales and symmetry 
of the cone, and compression is therefore an 
unlikely cause. Dimensions vary from about 
17 mm diameter and a length of up to 22 mm for 
the globose forms, to 15-22 mm diameter x 
20 mm in length for the ellipsoidal specimens. In 
apical view the pair of larger valves are at the 
ends of the apparent ellipse. The smaller, 
narrower, valves meet apically along a contact line 
of about 5 mm, thus separating the pair of larger 
valves (Fig. 4). Although apparently mature, the 
cones have a relatively smooth surface, with none 
of the roughness that characterises the opened 
cones of many Callitris species. However, one 
cone, at least, shows slight apically convergent 
ribs on the valves. This is reflected in an 
impression of what appears to be a decorticated 
cone (Fig. 3), having exposed ribs that have the 
same symmetry and relationships as the valves. 
None of the fruits has split to release its seeds, so 
the extent of the sutures separating the valves may 
not be fully expressed. Nevertheless, the sutures 
extend slightly more than halfway towards the 
base. The valves each have a near-apical, small 
(roughly 1 mm), circular to elliptical scar, 
arranged symmetrically on the fruit (Fig. 4). The 
origin or purpose of these scars is unclear, 
although they are in the same position as the 
'spurs' on the valves of some species of Callitris 
(Baker & Smith 1910: 47) and species of 
Papuacedrus, Widdringtonia and Tetraclinis 
(pers. obs.; Hill & Carpenter 1989; Mclver 2001). 
The base and attachment are seen in one 
specimen, showing a petiole expanding gradually 
to about 5 mm diameter at its junction with the 
fruit, where there is an expanded leaf scale below 
the smaller valve (Fig. 4). Another specimen 
seems to show a 7-8 mm diameter subcircular 
scar on the base, where the stem attached. 

Comparisons 

Initial comparisons were with species of 
Callitris. The obvious difference is in the number 



CUPRESSACEAN CONES IN THE EOCENE EYRE FORMATION 



187 




FIGURE 2. A. the fossil slab. SAM P 22732 B, positive cast of part of the slab, replicating the original appearance 
of the leaf litter at the time of burial. Scale in mm. 



188 



NS PLEDGE 



of valves in the cones, Callitris having three large 
valves alternating with three smaller ones, all 
ending acutely. Amongst living Australian 
conifers, there is only one species having two 
pairs of valves in the same decussate geometry as 
the fossils: Diselma archeri from Tasmania. 
However, its ovulate cones are tiny, barely larger 
in diameter than the shoots they terminate (pers. 
obs.). The living Papuacedrus from the island of 



New Guinea also has two pairs of valves, but its 
cones too are small and conical (Hill & Carpenter 
1989). 

A passing note in Baker and Smith (1910) 
directed attention to several non-Australian 
genera. Diselma had once been synonymised with 
Fitzroya from Chile and Patagonia, just as 
Callitris had been joined with Tetraclinis from 
Mediterranean North Africa and Widdringtonia 




FIGURE 3. Positive silicone rubber cast of the better fossil cones, showing 1. cone with apical view of sutures and 
valves with terminal scars; 2. cone with attached stem and basal leaf scale; 3, cone with basal attachment scar; 4, a 
decorticated cone; 5, foliage fragment. Scale in mm. 



CUPRESSACEAN CONES IN THE EOCENE EYRE FORMATION 



189 



spp. from southern Africa. 1 was not able to see 
fruits of Fitzroya, but am informed that they are 
nothing like the fossils (R. Hill, pers. comm., July 
2002), Through the help of Ms H. Vonow at (he 
South Australian Herbarium, T examined dry 
specimens of Diselma archeri, Tetraclinis 
artiatlata, Widdringtonia cupressoides, W. whytei 
and others and Papitacednis papuana, and living 
Widdringtonia spp. in the Mount Lofty Botanic 
Gardens. There is an obvious similarity between 



these species and the fossil cones in the number 
and symmetry of the valves. However, Tetraclinis 
can be removed from consideration on the basis 
of two characters; the spurs near the tips of its 
valves are quite enlarged, particularly on immature 
cones (there is obviously some allometric growth 
involved here), and the foliage structure is quite 
different from the fossil shoot associated with the 
cones. Offler (1969) indicates that Papuacedrus 
foliage is widely represented in her material, and 







FIGURE 4. Detail of cones; silicone rubber cast, showing 1, cone with apical view of sutures and valves with terminal 
scars. 2, cone with attached stem and basal leaf-scale, 3, cone with basal attachment scar, 4, foliage fragmenL 



190 



NS PLEDGE 



it has been recognised in the Eyre Formation of 
the Poole Creek Palaeochannel near Lake Eyre 
South (Alley & Pledge 2000: 69). Fossil foliage 
has also been recorded from various Oligocene 
and Miocene sites in Tasmania (Hill & Brodribb 
1999; Hill & Carpenter 1989). However, 
Papuacednis differs considerably from the fossils 
in the tear-drop shape and small size of the cones, 
and in the presence of bract-like appendages 
halfway up each cone scale (Hill & Carpenter 
1989). 

There are two genera considered closely related 
to Papuacednis, namely Libocedrus from New 
Zealand and New Caledonia, and fossil in 
Tasmania, and Austrocedrus from South America 
and also fossil in Tasmania (Hill & Carpenter 
1989). They have ovulate cones of the same order 
of size as Papuacedrus (and therefore are much 
smaller than the Lake Hart fossils) but are similar 
in having a structure subapically on the abaxial 
surface of each cone scale: a spine in the case of 
Libocedrus and a minute tubercle in Austrocedrus 
(ibid). However, the cone scales of Austrocedrus 
are all apically acute with none truncated; therefore, 
that genus can probably be ruled out of contention. 

The woody ovulate cones of the Widdringtonia 
spp. vary between species in both size and 
morphology. Like the fossils, they have two 
unequal pairs of valve scales, the upper pair 
having truncated apices that therefore meet along 
a contact of several millimetres. Of those 
examined, cones of W. cupressoides (Fig. 5) 
resemble the fossils most closely, although they 
tend to be smaller and have more prominent spurs 
on the valves, and the overall shape is more tulip- 
like, with a depressed base. Other species show a 
rougher, more rugose and warty or tuberculosc 
surface on the valves than do the fossils, but this 
may be a reflection of the maturity of the fruit. 
The Lake Hart fossils differ markedly from the 
North American Cretaceous species 
Widdringtonia americana (Mclver 2001), 
primarily in the development of a prominent umbo 
or boss on each valve scale in the latter species. 

The vegetative shoot (Figs 3, 4) associated with 
the fossil cones is poorly preserved (the grain-size 
of the sediment approaching that of the smaller 
features), and cannot be guaranteed to relate to 
the cones. It seems to have the same square cross- 
section and short decussate, scale-like leaves as 
Diselma arched (Clifford & Constantine 1980). 
This is unlike Widdringtonia cupressoides, which 
has shoots of rounder cross-section and longer 
decussate leaves. The leaves are apparently 
arranged in opposite pairs, as in Libocedrus and 




X1 




FIGURE 5. Female cones of Widdringtonia 
cupressoides (L.) Endlicher, specimens from the South 
Australian Herbarium collections, 

Papuacedrus (Hill & Carpenter 1989). 
Unfortunately, no other specimens from this site 
are available to support or deny this association, 
and the locality has reportedly been stripped by 
commercial interests for decorative stone. 



Conclusions 

It is apparent that these impressions represent 
the fruiting bodies of a previously unknown 
Australian species of conifer, and it is suggested 
that this shows closest affinities with species of 
Widdringtonia, Diselma and Papuacedrus. 
Without corroborative evidence in the form of 
organically associated foliage and pollen, it is not 
possible to specify which taxon the cones 
represent; no identification is proposed, but it 
could be a new extinct genus. Widdringtonia 
today inhabits seasonally dry to semi-arid regions 
of southern Africa (Hill & Brodribb 1999), as do 
Callitris spp. Such an environment is implied 
(Alley et al 1996) for part of the Eyre Formation. 
In view of the fact that Offler (1969) found no 



CUPRESSACEAN CONES IN THE EOCENE EYRE FORMATION 



»1 



trace of either Callitris or Actinosnvbtts foliage iii 
the 'silcrete flora' of the Woomera area, thai no 
cones of Callitris have subsequently been found 
in ihe large collections made since her study, and 
that there have been only a few finds of fossil 
Callitris foliage in southeastern Australia (Hill & 
Brodribb 1999), it is suggested that these typical 
Australian genera evolved subsequently, possibly 
from a form like the fossil taxon described Iterc. 



Acknowledgments 

I lhank Ms L Marsh and Mr T Nurenberg for 
donating the specimen to the South Australian Museum: 
Ms II Vonow. South Australian Herbal iuni. who 
assisted with access to preserved diy conifer specimens: 
and Prol. R Hill and a reviewer who read an early draft 
and provided some very useful references. 



RbHifiBNCF-S 



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Formation, lower Nelly Creek, southern Lake Eyre 
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CnllCOi RA & Cowley, WM. 1995. Billa Kalina Basin 
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Chapman, F. 1937. Descriptions of TcniaTy plant 
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Clifford, HT & Constantine, 1 1980. Ferns, Fern Allies 
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palacobotanical studies (exemplified by a study of 
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Adelaide. 

Offler, CE 1984 Extant and fossil Coniferales of 
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external morphology of the. vegetative shoots of the 
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Rowett, A, 1497 Earthwatch '96. MESA Journal 5: 27- 
29. 

Wopfner. H, Callen, RA & Hams, WK. 1974. The 
Lower TerUary Eyre Formation of the southwestern 
Great Artesian Basin. Journal of the Geologii ol 
Socun of Australia. 2I( I): 17-51. 



DESCRIPTIONS OF A NEW GENUS AND TWO NEW SPECIES OF 

VIVIPARID SNAILS (MOLLUSCA: GASTROPODA: VIVIPARIDAE) FROM 

THE EARLY CRETACEOUS (MIDDLE-LATE ALBIAN) GRIMAN CREEK 

FORMATION OF LIGHTNING RIDGE, NORTHERN NEW SOUTH WALES 

Robert J. Hamilton-Bruce, Brian J. Smith & Karen L. Gowleti -Holmes 

Summary 

Fossil gastropods belonging to the family Viviparidae (Caenogastropoda (Prosobranchia)) are 
described from the Early Cretaceous (middle-late Albian) Griman Creek Formation of Lightning 
Ridge, northern New South Wales. From our research, this is the earliest definitive record of non- 
marine gastropods from Australia and among the oldest viviparid material in the world recorded to 
date. A new genus and two new species are described, and other material attributable to the extant 
genus Notopal is recorded. Implications for the current distribution of the Viviparidae are 
addressed. 



DESCRIPTIONS OF A NEW GEMS AND TWO NEW SPECIES OF VIVIPARID SNAILS 
(MOLLUSCA: GASTROPODA: VIVIPARIDAE) FROM THE EARLY CRETACEOUS 
(MIDDLE-LATE AI.BIAN) GRIMAN CREEK FORMATION OF LIGHTNING RIDGE, 

NORTHERN NEW SOUTH WALES 



ROBERT J HAMILTON-BRUCE. BRIAN J SMITH & KAREN L GOWLETT-HOLMES 



HAMILTON BRUCE. RJ. SMITH. BJ & OOWLETT-HOLMES. KL. 2002. Descriptions of a 
How genus and two new species of viviparid snails (Mollusea: Gastropoda: Vivip;ai((ae) from 
the Early Cretaceous (middle- lute Albianl Griman Creek Formation of Lightning Ridge, 
northern New South Wales. Records of ihe South Australian Museum 35(2): 193-203. 

Fossil gastropods belonging to the family Viviparidae (Caenogastropodn tProsobranehuO) 
are described from Hie Early Cretaceous (middle- late Albian) Griman Creek Formation of 
Lightning Ridge, northern New South Wales From our research, this is the earliest definitive 
record of non-marine gastropods from Australia and amongst the oldest viviparid material in 
'he world recorded to dale. A new genus and two new species are described, and other material 
jiiiihniaiile 10 |de extant genus Nniopala is recorded. Implications for the current distribution 
of the Viviparidae aie addressed, 

"•Robert .1 Hamilton-Bruce, South Australian Museum, North Terrace, Adelaide,. South 
Australia 5000; Brian J Smith. Queen Victoria Museum. Wellington Street, Launeeston. 
Tasmania 7250, Australia; Karen L Gowlctt-Holmes, CSIRO Division of Marine Research, 
GPO Bo\ 1538, Hobart, Tasmania 7001. Australia; ^corresponding author — email: Hamilton 
bruce.rohertts'saugov.sa. gov.au Manuscript received 29 July 2002. 



Opalised fossils have long been known from 
the Lower Cretaceous (middle-laie Albian) 
freshwater deposits of the Griman Creek 
Formation, Lightning Ridge, New South Wales 
(Smith 1999). To date, much of the published 
work has focused on Fish (Kemp 1991; Kemp & 
Molnar 1981) and terrestrial tehapod remains 
(Archer ct al 1985; Flannery et a| 1995; Molnar 
I'WJa, 1980b, 1991, Molnar & Gallon 1986; Rich 
et al 1989) with relatively few reports (Detimau et 
al 1992; McMichael 1957; Smith 1999) 
summarising the vast quantities of invertebrate 
material recovered from the area. The recent 
opportunity to examine Lightning Ridee 
gastropod specimens (brought to our attention by 
Ben Kear. South Australian Museum, as part of 
an ongoing assessment of vertebrate and 
invertebrate fossil material from the locality) from 
the Collections ol the Australian Museum. Sydney, 
and several private individuals has prompted a 
systematic appraisal of the gasiropods within ihe 
material recovered. This study describes 
specimens attributable to the family Viviparidae 
(Caenogastropoda (= Prosobranchta)), including a 
new genus and two new species. Indeterminate 
material belonging to the exiant genus Noiopala 
is also recorded, and the implications for 



distribution of extant viviparid taxa are discussed. 

The Viviparidae is a cosmopolitan group of 
freshwater caenogasiropods, characterised by 
medium- to large-si«:d lurbiniform shells which 
possess a rounded hody whorl, moderately high, 
pointed spire; wide, round aperture; and sub- 
spiral, horny operculum (Smith 1992). The current 
Australian distribution of the group is limited to a 
few species occurring in lite large drainage basins 
that span much of ihe arid centre and northern 
tropical regions of the continent 

The fossil record for Viviparidae is known from 
the Jurassic to Recent {Vniparus Monitor!, 1810), 
with a tentative report based on an internal shell 
(Wemicia Cox, 1927) mould (possibly of marine 
origin) from the Lower Carboniferous of England 
(Brookes-Knight et al 1960). Within Australia 
there are few records of fossil viviparids. 
Elheridge (1902) described the earliest potential 
Australian taxon, VMporus (?) atba-scopularis 
Elhendge (also noted by Newton (1915)) from the 
Aprian marine deposits of Ihe Doncastcr Member 
(Wallumbilla Formation \ensu Burton & Mason 
I998), White Cliffs, New South Wales This 
specimen is currently under examination by the 
authors and at present is regarded as being of 
unclear viviparid affinity. Cotton (1935a) erected 



194 



RJ HAMILTON-BRUCE, BJ SMITH, KL GOWLETT-HOLMES 



a species of Notopala (N. wanjacalda) from late 
Pleistocene sediments along the Murray River 
near Sunnyside, South Australia, and also noted a 
second taxon (Notopala sp.) from the same 
deposits, which showed strong similarity to the 
extant N. hanleyi (Frauenfeld, 1862). Viviparid 
snail shells from Early Cretaceous deposits in the 
Lightning Ridge area were recorded but not 
described by Dettman et al (1992) and Smith 
(1999); the latter also recorded possible 
representatives of the Naticidae, Thiaridae and 
Ellobiidae. Few other Australian non-marine 
gastropod fossils (all of Tertiary age) have been 
recorded (Archer et al 1994; Arena, 1997; 
Chapman 1937; McMichael 1968). 



Materials and Methods 

All specimens described herein are derived 
from the Lightning Ridge opal fields (exact mine 
localities from which these specimens originated 
are unknown), Surat Basin, northwestern New 
South Wales. The opal-bearing sediments in this 
area form part of the Griman Creek Formation, a 
unit dated as middle-late Albian in age (Dettman 
et al 1992). The deposit predominantly reflects a 
coastal plain facies (Burger 1988; Dettman et al 
1992) situated in an Early Cretaceous high latitude 
zone (-70° S, Embleton 1984). Invertebrate 
fossils, plant root impressions and vertebrate 
taphonomy suggest a freshwater estuarine to 
lacustrine setting (Dettman et al 1992; Molnar 
1980a). Palaeoclimatic indicators imply strongly 
seasonal conditions with sea-level isotopic 
palaeotemperatures in the Eromanga Basin / Surat 
Basin regions ranging from 11.9 C C (northeast) to 
16.3°C (southwest) (Dettman et al 1992; Stevens 
& Clayton 1971). 

Designation of parietal fold position on the 
specimens was achieved by positioning the fold 
within the aperture on a 360° compass setting 
when a vertical line through the axis, juxtaposed 
to the column, is intersected by a horizontal line 
corresponding to the midpoint of the aperture. 
The adapical axis above the point of intersection 
is taken as zero degrees. The diameters of all 
shells were measured following the method of 
Boycott (1928) and are defined as 'the greatest 
dimension that can be found starting with the 
edge of the lip to a point on the opposite side of 
the shell on the last whorl'. To enable 
extrapolation of the numbers of missing whorls, 
the incremental angle of the shells was judged 
by drawing a line so as to touch each of the 



present whorls across each side of the shell; the 
point at which they met was assessed as being 
approximately the original apex of the shell (Cox 
1960). All specimens are deposited in the 
Australian Museum (AM), Sydney. All 
specimens were prepared using a diamond drill- 
bit attached via a flexible lead to a high-speed 
rotary motor. Shell measurements were made to 
the nearest 0.05 mm using dial calipers. 



System atics 
Class GASTROPODA 
Subclass ORTHOGASTROPODA 
Superorder CAENOGASTROPODA 
Order ARCfflTAENIOGLOSSA 
Superfamily AMPULLARIOIDEA 
Family VIVIPARID AE Gray, 1847 



Diagnosis 

Medium to large dextral, turbiniform shells, 
body whorl rounded, spire moderately high, 
pointed; aperture wide, round, parietal fold 
present or absent; operculum horny, subspiral 
(modified after Smith 1992). 

Remarks 

The above diagnosis follows Smith (1992), 
modified to accommodate the presence of a 
parietal fold in the new genus described below. 
Viviparid snails are, as their name suggests, 
viviparous (live bearing) and are found in both 
lotic and lentic systems throughout the world 
(Browne 1978). Within Australia the family is 
currently represented by the extant native genera 
Notopala, Larina and Centropala (Smith 1992). 
A species accidentally introduced from Asia, 
Bellamya heudei guangdungensis (Kobelt, 1906), 
recorded by Shea (1994) as established in New 
South Wales, is not considered part of the 
Australian fauna in this study. The Australian 
members of the family have undergone substantial 
revision over many years (Cotton 1935a, 1935b; 
Sheldon & Walker 1993; Stoddart 1982); the use 
of intraspecific shell variation and morphometric 
data by recent authors has resulted in a substantial 
reduction in the numbers of accepted species. 
Shell colouring and pattern and the form of the 
operculum, which are important characters in 



EARLY CRETACEOUS VIVIPARID SNAIt.S 



195 



determining generic placement in this family, arc 
characters lost in most fossils. 

The present specimens from Lightning Ridge 
are over 100 million years old (Albian [108- 
97.5 MaJ) and totally opalised. While sculpturing 
on some specimens has been preserved, colour- 
bands and opcreula have not; therefore, only 
Structural shell characters could be used to assign 
them to taxa. Two major factors have been taken 
into account before deciding to place them in 
Vivipandae — the shell morphology, which fits 
within the currently accepted diagnosis of the 
family widi only slight modification (the presence 
of the parietal fold in Ihe two new species); and 
the freshwater depositional environment from 
which they originated. Whilst all the specimens 
can be accommodated within the family 
Viviparidae, those possessing a parietal fold 
cannot be assigned to any currently recognised 
genus and are considered to belong to a new 
genus which is described below. Direct 
comparisons between species, living or fossil, are 
few (Brown 1980); however, due to the lack of 
colour patterns and opcreula as well as soft parts 
in these fossils, the only remaining methods are 
based on morphology. Therefore, we have applied 
parts of the morphometric data gathered by 
Sheldon and Walker (1993) to justify the 
placement of these specimens within Viviparidae, 
and in one case in the extant genus Notopala, and 
to offer a method of comparison between extant 
and extinct species. None of the data comparisons 



are in any way intended to suggest relationships 
between members of either the Notopala sp. 
presented here or the new genus and any extant 
species. 



Genus Albianopalin gen. no v. 

Diagnosis 

Shell dextral, thick, solid, globose to 
subglobose, ventricose, three-five whorls, 
subumbilicale; aperture subovate, large, 
approximately cqua] to height of spire; parietal 
fold present, simple. Operculum unknown. 

Type species 
Albianopalin benkeari sp. nov. 

Etymology 

From the combination of Albian, n. referring to 
age of the Lower Cretaceous opal-bearing deposits 
of the Griman Creek Formation, and opalin. 
Middle English, from apalus, Latin, alteration of 
Greek opallios, for 'opal'. 

Remarks 

While this new genus has similarities to 
Notopala (Cotton, 1935b), Albianopalin gen. nov. 
is easily separated from Notopala and all other 
genera within the family by the presence of a 
parietal fold in the basal lip, currently a feature 
unique to this genus within the family. 





B 



RCil.JRE 1 . AMFI22185 Albaiw/uilin benkeari sp. nov. in A, apertural and H. dorsal views. 



196 



RJ HAMILTON-BRUCE, BJ SMITH, KL GOWLETT-HOLMES 




Etymology 

Named for Ben Kear, 
study. 



for his assistance in this 



FIGURE 2. AMF122185 Albanopalin benkeari sp. nov. 
in apertural view showing angle of parietal fold. 



Albianopalin benkeari sp. nov. (Figs 1, 2, 
Appendix) 

Diagnosis 

With the features of the genus; parietal fold 
simple, 70° from vertical axis. 

Type specimen 
' Holotype: AMF122185 (Figs 1, 2, Appendix) 

Locality and horizon 

Lightning Ridge opal fields (precise site within 
the opal fields unknown), northern New South 
Wales, Surat Basin, Griman Creek Formation, 
middle-late Albian. 

Description 

Shell 17.7 mm high, 16.4 mm maximum 
diameter, dextral, turbiniform, subglobose. 
Teleconch with three complete whorls and broken 
parts, pointing to the possible presence of further 
whorls. Whorls impressed. Relatively evenly 
spaced spiral prosocyrt ornamentation present on 
many areas of teleconch. Aperture large 
(10.85 mm high), round; basal outer lip varicose 
with no evidence of eversion. A single, 
triangulated parietal fold (3.0 mm long, 1 .0 mm 
wide, 2.0 mm deep) present on upper surface of 
basal lip, deepest side facing adaperturally (Fig. 
2). 



Remarks 

Holotype unique. The position of the parietal 
fold separates A. benkeari sp. nov. from the 
other new species described below. The 
holotype specimen is undistorted, but the spire 
is incomplete. Extrapolation from the spire 
angle indicates that there may have been at least 
four and as many as five complete whorls 
originally. Being opalised, the state of 
preservation of the holotype is very good, with 
much of its detailed surface ornamentation 
preserved. 



Albianopalin Uzsmithae sp. nov. (Figs 3, 4, 
Appendix) 

Diagnosis 

With the features of the genus; parietal fold 
simple, 1 12° from the vertical. 

Type specimen 
Holotype: AMF122186 (Figs 3, 4, Appendix) 

Locality and horizon 

Lightning Ridge opal fields (precise site within 
the opal fields unknown), northern New South 
Wales, Surat Basin, Griman Creek Formation, 
middle-late Albian. 

Description 

Shell, 15.1 mm high, 12.9 mm maximum 
diameter, dextral, turbiniform, subglobose. 
Teleconch with two complete whorls and broken 
parts, pointing to the possible presence of further 
whorls. Whorls impressed. This particular 
specimen has undergone some dorsoventral 
compression which has resulted in distortion, 
especially at base of final whorl. Aperture large 
(9.95 mm high), round; basal outer lip varicose 
with no evidence of eversion. A single, basal 
parietal fold (3.75 mm long, 2.3 mm wide, 
2.2 mm deep) is present on surface of basal lip, 
deepest side facing adaperturally (Fig. 4). Spiral 
prosocyrt ornamentation present on parts of 
teleconch, especially proximal to aperture. 

Etymology 

We name this species for our colleague 
Elizabeth (Liz) Smith of the Australian Museum 
(AM), for her assistance in this study. 



EARLY CRETACEOUS VIV1PARID SNAILS 



197 





HH 



A B 

FIGURE 3. AMF1 22186 Albunnpulin lizsmiilwe sp, nov. in A, apertural and B, dorsal views. 




FIGURE 4. AMFI22186 Albanopalin Uzsmithae sp. 
nov. in apcrlural view showing angle of parietal fold. 



Remarks 

Holotype unique. The position of the parietal 
fold separates A. Uzsmithae sp. nov. from A. 
benkearl sp. nov. as described above. The 
holotype has undergone moderate dorsoventral 
compression, resulting in the specimen appearing 
stretched. Like A. benkeari sp. nov., the spire is 
incomplete, and extrapolation from [he spire angle 
indicates that there originally may have been at 
least four and as many as five complete whorls. 
Being opalised and having damage to parts of the 
surface of the teleconch, the state of preservation 
is good enough to preserve some of its detailed 
surface ornamentation. 



Genus Notopala (Cotton, 1935b) 



Type specimen 

Type species: Paludina hanleyi (Frauenfeld. 
1864) by original designation. 

Diagnosis 

Shell dextral, globose-conic, subumbilicaic, 
five whorls, ventricosc to angulate below the 
periphery; aperture subovate, large, about equal to 
height of spire; parietal fold absent; operculum 
corneous. 



198 



RJ HAMILTON-BRUCE, BJ SMITH, KL GOWLETT-HOLMES 





B 



FIGURE 5. AMF13001 Notopala sp. in A, apertural and B, dorsal views. 



Remarks 

These specimens are extremely significant. 
Until their discovery, none of the currently living 
native Australian viviparid genera were known 
from deposits older than Plio-Pleistocene. This 
temporal range is now extended back to the Lower 
Cretaceous (middle-late Albian). It is also 
interesting to note that another living species, the 
lungfish Neoceratodus forsteri, has been recorded 
from the Griman Creek Formation of Lightning 
Ridge (Kemp & Molnar 1981). 

Notopala sp. (Fig. 5, Appendix) 

Referred material 

AMF13001 (Fig. 5), AMF122166-AMF122184 
(not figured herein). 

Locality and horizon 

Lightning Ridge opal fields (precise site within 
the opal fields unknown), northern New South 
Wales, Surat Basin, Griman Creek Formation, 
middle-late Albian. 

Description 

Shell dextral, turbiniform, subglobose. 
Teleconch with two to three complete 
impressed whorls; the incremental angle of the 
shell indicates that there were originally four or 
five whorls. Aperture large, round; basal outer 



lip varicose with no evidence of eversion. 

Remarks 

There is considerable variation in size, state of 
preservation, degree of corrosion and/or damage to 
the shells, opalisation, amount of matrix deposition 
on the shells, completeness of the aperture and 
presence/absence of ornamentation. We have not 
described these specimens as a new species at this 
stage. We feel that the exact taxonomic position of 
this material compared with other species in the 
genus requires further study. The preceding table 
(Table 1), diagram (Fig. 6, modified from Sheldon 
& Walker 1993) and graph (Fig. 7) show 
measurements of shell characters for each of the 
living Australian species of Notopala, and illustrate 
morphometric similarity of the specimens of 
Albianopalin nov. sp. as well as Notopala sp. 
described here to existing members of the genus. 
However, the specimens of Notopala sp. vary 
greatly in their physical condition, and it has not 
been possible to take the full range of 
measurements from each of the 20 specimens being 
studied. The reason for the inclusion of this data is 
not to demonstrate any relationship between extant 
species and the specimens from Lightning Ridge, 
but to show that the physical characteristics of the 
new material {Albianopalin sp. nov. and Notopala 
sp.) fit within the parameters for inclusion in the 
family Viviparidae. 



EARLY CRETACEOUS VIVIPARID SNAILS 



199 



Aperture Lip 
tcngth 
(ALL) 
Umbilical "^ 
widih 

CVMB) 




Shell 
length 
(SHL) 



FIGURE 6. Diagram illustrating measurement parameters used in the morphomctric analysis (modified from 
Sheldon & Walker 1993). 



APL 



SHW UMB 



APW 



ALL 




>:•:•>: 

'.•$. 



CH Notopala sp.. gen. 
' "Banded" Shells 

Notopala sublineata (Cooper) 
I Notopala sublineata (Darling) 
N. hanleyi 

I = Standard deviation 



N. waterhousii 

N. essingtonensis 

N. alisoni 

A. benkeari nov, sp, * 

A. lizsmithae nov. sp. * 

* single sample 



FIGURE 7. Histogram showing results of morphometric analysis. Taxa include living species of Notopala (modified 
from Sheldon & Walker 1993), Notopala sp. (AMF1300I, AMF122185-122184), Alhianopalin benkeari 
(AMF122185) and A. lizsmithae (AMF122186). 



200 



RJ HAMILTON-BRUCE, BJ SMITH. KL GOWLETT-HOLMES 



TABLE 1. Mean shell measurements (mm) for various living species of Notopala (*modified from Sheldon & 
Walker 1993) compared with those for Notopala sp. (AMF13001, AMF122185-122184), Albumopalin benkeari 
(AMF122185) and A. lizsmithae (AMF122186). Because not all specimens of Notopala sp. were complete, the 
number (n) of specimens used for each individual parameter is indicated separately. Standard deviation (SD) rows 
were not included for A. benkeari and A. lizsmitliae as only a single specimen has currently been recovered for each 
of these taxa. Abbreviations: SHL, shell length; APL, apertural length; SHW, shell width; UMB, umbilical width; 
APW, aperture width; SPL, spire length; ALL, aperture lip length. 



Species 


Number 


SHL 


APL 


SHW 


UMB 


APW 


SPL 


ALL 


*N. alisoni 


20 




21.59 


11.91 


16.3 


02.59 


09.77 


14.78 


05.35 


(Brazier, 1979) 




SD 


0.086 


0.039 


0.054 


0.014 


0.034 


0.064 


0.015 


*N. essingionensis 


20 




21.70 


12.00 


16.58 


1.96 


10.10 


14.47 


6.11 


(Frauenfeld, 1862) 




SD 


0.082 


0.038 


0.067 


0.016 


0.038 


0.067 


0.025 


*N. waterhousii 


10 




38.59 


22.84 


33.14 


4.63 


18.88 


25.01 


10.02 


(Adams & Angus, 1864) 




SD 


0.209 


0.105 


0.164 


0.024 


0.087 


0.131 


0.043 


*N. hanleyi 


42 




16.67 


10.50 


13.83 


1.29 


8.52 


11.00 


5.57 


(Frauenfeld, 1862) 




SD 


0.520 


0.026 


0.036 


0.00 


0.025 


0.037 


0.013 


*N. sublineata 


28 




18.32 


10.84 


14.53 


2.05 


9.21 


12.04 


5.24 


(D) (Conrad 1850) 




SD 


0.063 


0.024 


0.040 


0.012 


0.026 


0.039 


0.016 


*N. sublineata 


35 




19.46 


11.06 


15.52 


2.21 


8.89 


12.59 


4.93 


(C) (Conrad 1850) 




SD 


0.058 


0.027 


0.039 


0.010 


0.018 


0.040 


0.009 


♦'Banded' shells 


15 




22.15 


13.18 


18.10 


2.06 


10.69 


14.6 


6.42 






SD 


0.110 


0.053 


0.008 


0.011 


0.045 


0.079 


0.029 


Notopala sp. 


20 




- 


10.35 (19) 


14.43 (20) 


3.24(10) 


9.20 (15) 


- 


4.71 (9) 






SD 


- 


2.563 


3.053 


0.616 


1.80 


- 


1.071 


A. benkeari 


1 




17.7 


10.85 


16.4 


3.35 


9.85 


— 


6.2 


A. lizsmithae 


1 




15.1 


9.95 


12.9 


3.4 


8.15 


- 


4.4 



Discussion 

Albianopalin benkeari sp. nov., A. lizsmithae 
sp. nov. and Notopala sp. share the distinction of 
being the three oldest known definitively assigned 
members of the Viviparidae in Australia and thus 
serve to extend the range of the family in this 
region back to at least the uppermost Early 
Cretaceous. Since the family is confidently known 
from the Jurassic to Recent in Europe, Viviparidae 
therefore appears to be an ancient pre-Jurassic 
group of probable Pangean origin. By the 
Cretaceous, the family had diversified within the 
Gondwanan region into a range of endemic genera 
and species. 

While there are numerous Cretaceous records 
of freshwater bivalves from Australia (Dettman et 
al 1992; Hocknull 1997; Jell & Duncan 1986; 
Ludbrook 1985; McMichael 1957), there are very 
few for non-marine gastropods from the same 
period. The reasons for this apparent absence are 
unknown, but could be related to preservational 
biases (with shells rapidly breaking up or 
dissolving after death). Another possibly reason 
could be a lack of gastropod species due to poor 
tolerance of the strongly seasonal near-freezing 



climatic conditions which characterised many 
Australian high latitude depositional environments 
during the Early Cretaceous. 

The implications of this study, combined with 
subsequent investigations, could eventually 
demonstrate a Gondwanan and possibly 
Australian radiation for many of the species living 
around the globe today. 

Acknowledgments 

This paper would never have been initiated without 
Ben Kear, who, since its inception, has more than 
earned our sincere thanks. We wish also to thank Philip 
Ryan for examining the statistical methods employed. 
Greg Brown and George Koulis of the Royal Adelaide 
Hospital (RAH) helped considerably by contributing 
high-grade imaging. Elizabeth (Liz) Smith. Bob Jones, 
Henk Godthelp, Tom Rich, Rolf Schmidt and Neville 
Pledge supported this work and generously permitted 
access to their collections. Philip Clarke of the South 
Australian Museum, Adelaide (SAMA), gave financial 
support and encouragement. Anne Hamilton-Bruce of 
The Queen Elizabeth Hospital (TQEH) contributed 
lateral thinking and assistance with editing. Chris lzzo 
(zoology student at Adelaide University) assisted with 
measuring and statistical data analysis. Origin Energy, 



EARLY CRETACEOUS VIVIPARID SNAILS 



201 



The Advertiser newspaper. Coober Pcdy Tom ism 
Association, llie Souih Australian Museum and the 
Walcrhousc Club gave other support to this project. 



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EARLY CRETACEOUS VIVIPARID SNAILS 



203 



APPENDIX 



Measurements (mm) of all Lightning Ridge gastropod specimens used in this study (AMF13001, 
AMF122166-AMF122186). Abbreviations: SHL, shell length; APL, apertural length; SHW, shell 
width; 1MB, umbilical width; APW, aperture width; SPL, spire length; ALL, aperture lip length. Spire 
length measurements are omitted owing to incomplete apices in all specimens sampled. 



Specimen 




SHL 


APL 


SHW 


UMB 


APW 


ALL 


1. AMF 122 185 (type A. 


benkeari) 


17.70 


10.85 


16.40 


3.35 


9.85 


6.20 


2. AMF 1221 86 (type A 


lizsmithae) 


15.10 


9.95 


12.90 


3.40 


8.15 


4.40 


3. AMF13001 (Notopala sp.) 


19.25 


12.00 


17.30 


3.75 


11.10 


5.02 


4. AMF122166 




17.65 


11.05 


14.50 


- 


8.55 


— 


5. AMF122167 




14.37 


8.75 


12.75 


8.50 


— 


_ 


6. AMF122168 




11.60 


- 


11.85 


2.30 


7.80 


3.35 


7. AMF122169 




15.62 


8.80 


12.45 


3.00 


— 


5.00 


8. AMF1 22170 




12.60 


8.80 


11.65 


3.00 


7.65 


4.35 


9. AMF122171 




19.00 


13.25 


19.90 


- 


12.20 


- 


10. AMF122172 




20.42 


12.10 


17.20 


4.20 


- 


6.00 


II. AMF122173 




18.00 


11.05 


16.20 


3.90 


9.80 


_ 


12. AMF122174 




17.92 


11.20 


15.55 


- 


11.15 


_ 


13. AMF1 22175 




19.82 


12.63 


17.85 


3.35 


— 


_ 


14. AMF122176 




19.35 


11.10 


16.75 


- 


10.50 


_ 


15. AMF122177 




19.57 


12.65 


17.50 


— 


11.70 


_ 


16. AMF122178 




18.55 


10.05 


15.15 


- 


- 


— 


17. AMF122179 




17.95 


11.30 


15.20 


3.80 


9.60 


4.80 


18. AMF122180 




10.22 


6.50 


9.00 


- 


_ 


6.00 


19. AMF122181 




6.92 


7.20 


9.40 


2.60 


6.15 


3.60 


20. AMF 122 182 




9.32 


6.50 


8.75 


2.35 


5.60 


3.15 


2LAMF122183 




17.80 


11.70 


16.00 


- 


9.80 


_ 


22. AMF 122 184 




- 


16.10 


10.10 


13.40 


- 


8.30 



SOUTH 

AUSTRALIAN 

MUSEUM 



VOLUME 35 PART 2 
OCTOBER 2002 
ISSN 0376—2750 



CONTENTS: 



ARTICLES 

105 CHS WATTS 

The larvae of some Australian aquatic Hyclrophilidae 



otera: Insecta) 



39 AM PINDER & PK McEVOY 

Embolocephaltis yamaguchu (Brinkhurst, 19" 



idae) from South 



147 PA CLARKE 

Early Aboriginal 



he Lower 



169 MN HUTCHINSON & BS MACKNESS 



Queensland, with a descriptii 



new speeies 



185 NS PLEDGE 
First report ol 



s in the Eocene Eyre Formation of South Australia. 



93 RJ HAMILTON-BRUCE, BJ SMITH & K.L GOWLETT-HOLMES 

Descriptions of a new genus and two new species of viviparid snails (Mollusca: 
Gastropoda: Viviparidae) from the Early Cretaceous (middle — late Albian) Griman Creek 
Formation of Liehtnine Ridue, northern New South Wales. 



Published by the South Australian Museum. 
North Terrace, Adelaide, South Australia 5000.